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ISSN 0374-7859 


THE GARDENS’ BULLETIN 
SINGAPORE 


Volume 38 
(1985) 


A periodical reflecting the interests and 
activities of the Botanic Gardens 
Singapore 


Published by the Botanic Gardens 

Parks and Recreation Department 

Ministry of National Development 
Cluny Road, Singapore 1025. 


Printed by Eurasia Press 


CONTENTS 


Volume 38 
PART 1 — Ist June 1985 Pages 
CORNER,.E.J.H: 
waeettowny Gr seme Isicts Fast of Pahane afd JOROTE ooo. 5... 12. on 5 ence ce ewe sneecseesnbessenceenees 1-42 


HOTTA, Mitsuru: 


New Species of the Genus Homalomena (Araceae) from Sumatra with a Short Note on the 
ieee PREMARIN © 2 PER SOM re eg al ol be ni sn eRe Wd ad PSD TET,“ GE ND, Fe AS 43-54 


WILDE, W.J.J.O. DE: 
A New Account of the Genus Horsfieldia (Myristicaceae), Pt 2 ..............c cece eee eee eee ee nes 55-144 


PART 2 — Ist December 1985 


HOLTTUM, R.E.: 


Two New Species of Tectaria from Limestone in Peninsular Malaysia, with Comments 
OO OS Se eee on RRORE SER 50 oe La or: | (need a ne 145-148 


KENG, Hsuan: 
Seen ee hE PERE E TANANS (ESM AROSE (0G) ) 25. ores oe cs detec Leela das. ed vagecacabeccees en 149-174 


LIM-HO CuHEE LEN and LEE SING Kona: 


Micropropagation of Lagerstroemia speciosa (L.) Pers. .............ecececececenenceceeeeeceneneeeeenees 175-184 


WILDE, W.J.J.O. DE: 
A New Account of the Genus Horsfieldia (Myristicaceae), Pt 3 .............. 0. ccc ececeeeeeeeeee ences 185-225 


BIDIN, Aziz, and TREvoR WALKER: 


Comparative Anatomy of the Stipe of the Fern Genus Adiantum L. ...................00000e0e0e02+. 227-233 


gE GES ae at, pe te ee gt ge oe es 8 oc. 2 a 235-242 


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INDEX 
Volume 38 


Basionyms and synonyms appear in italics. Page numbers in italics indicate the presence of illustra- 
tions. ‘Vernacular Names’ is a new entry. 


Acanthospermum australe 167 Artemisia lactiflora 168 
brasilum 167 vulgaris 168 
Acrostichum aureum 10 Arthrophyllum ovalifolium 17 
Adenanthera pavonina 4, 9, 17 Aspidium 147 
Adenosacme malayana 159 Asplenium glaucophyllum 25 
Adenostemma lavenia 168 macrophyllum 25 
viscosum 168 nidus-avis 12, 24 
Adiantum see also under ‘Anatomical structure Atalantia 12, 14 
< Atalantia forest 36 
Adiantum stenochlamys 25 Atalantia monophylla 6, 9, 17, 35 
Adina rubescens 150 Atalantia wood 6, 8, 9 
Aganosma 9 Baeckia 16 
marginata 10, 21 Barringtonia 39 
Agathis forest 94, 106, 186 asiatica 17 
Agathis-Casuarina forest 224 macrostachya 14, 17 
Ageratum conyzoides 168 Bidens pilosa 168 
Aidia cochinchinensis 162 Blechnum orientale 25 
corymbosa 162 Blumea balsamifera 168 
Allophyllus cobbe 8, /0, 14 lacera 168 
var. glaber 14 Borreria alata 150 
var. limosus 14, 17 articularis 150 
var. marinus 11, 14, 17 hispida 150 
var. velutinus 14, 17 laevicaulis 150 
javanicus 14 latifolia 150 
longipes 14 setidens 150 
racemosus 14 Breynia coronata 14 
ternatus 14 Bruguiera gymnorrhiza 4, 5 
timorensis 14 Bucephalandra 43 
Anatomical structure of stipes in ferns 227 Buchanania arborescens 17 
in Adiantum 228 Bulbophyllum vaginatum 23 
formosum 232 Caelospermum 151 
species list 230, 231 Calamus burkillianus 15 
stelar types 228 chibehensis 6, 7, 15, 21 
xylem shape in: Callicarpa longifolia 11, 17 
Adiantum capellis-veneris 232 Calophyllum-Ficus forest 67 
A. Capellis-Veneris Group 232 Canavalia turgida 23 
A Caudatum and Reniform Groups 228 Cansjera zizyphoides 23 
A. formosum 232 Canthium confertum 150 
A. hispidulum, cunninghamii, affine, dicoccum 150 
whitianum, sylvaticum & pedatum 232 glabrum 150 
A. lucidum and tetraphyllum 228 horridum 150 
A. lunulatum 228 molle 150 
A. macrophyllum & serratocristatum 232 Caprifoliaceae, key to genera 166 
A. patens 232 Caryota mitis 6, 17 
A. reniforme 232 Cassytha filiformis 22 
A. tetraphyllum 232 Castanopsis forest 86, 94 
illustrated 229 Castanopsis-dominated forest 67 
Aneilema 3 Castanopsis-dominated ridge 111 
Aneilema sp. 23 Castanopsis-Lithocarpus forest 134 
Anisoptera-Hopea-dominated forest 111 Casuarina 3 
Anisoptera-mixed forest 122 equisetifolia 14, 17 
Antidesma cuspidatum 17 Cedrela 17 
Aporosa? 15 Celastrus 15, 17 
Ardisia crispa 17 Centipeda minima 168 
elliptica 17 Cephaelis singaporensis 151 
Aridarum 43 Cerbera manghas 17 


235 


236 Gard. Bull. Sing. 38 (1985) 


Chassalia 151 Diplospora malaccensis 18, 152 
chartacea 151 Dipterocarp forest 209, 217 
curviflora 23, 151 Dischidia rafflesiana 24 
pubescens 151 Dracaena maingayi 18 

Chionanthus ramiflorus 17 Drynaria quercifolia 24 

Chrysanthemum indicum 168 Dryobalanops forest 209 
morifolium 168 Dryobalanops fusca-dominant forest 215 

Chrysopogon 12 Dryopteris glabrior 148 
fulvus 11, 15, 38 Eclipta alba 169 

Cissampelos? 14 prostrata 169 

Cissampelos sp. 4 Ehretia 22 

Cissus repens 22 Ehretia sp. 15 

Clerodendron inerme 4 Elaeocarpus floribundus 18 

Cocculus ovalifolius 17 Elatostema 4 

Coelorrachis glandulosa 23 Elatostema sp. 4 

Coelospermum scandens 151 Elephantopus scaber 169 

Coffea canephora var. 151 Eleutheranthera ruderalis 169 
liberica 151 Emilia sonchifolia 169 
robusta 151 Endocomia 75 

Colubrina asiatica 4 macrocoma 106 

Commersonia platyphylla 23 rufirachis 210 

Compositae, key to genera 167 Enydra fluctuans 170 

Connarus monospermus var. malayana 22 Erechtites hieracifolia 170 

Coptosapelta flavescens 152 valerianifolia 170 
griffith 151 Erigeron sumatrense 170 
parviflora 151 Erioglossum rubiginosum 18 
tomentosa 152 Erythrina indica 18 

Cordia subcordata 18 Erythroxylon cuneatum 18 

Coreopsis lanceolata 169 Eucalyptopsis-dominated forest 94 
tinctoria 169 Eculina longiflora 162 

Cosmos sulphureus 169 Eugenia claviflora 18 

Crassocephalum crepidioides 169 glauca 18 

Crepis japonica 174 grandis 2, 3,5, 6,6, 7,9, 11,12; ty Eee 

Croton heterocarpus 18 Eugenia grandis forest 12 

Ctenitopsis 146 Eugenia grandis zone 4 

Cycas 2,4 longiflora 18 
rumphii 18 palembanica 18 

Cyclophorus adnascens 25 polita 18 

Cymbidium finlaysonianum 24 subdecussata 18 

Cynometra ramiflora 18 Eulalia ridleyi 23 

Cyperus cyperinus 3, 23 Eupatorium odoratum 170 
diffusus 23 Eurycoma longifolia 6, 18 
dubius 3, 23 Excoecaria agallocha 18 
javanicus 3, 6, 7, 10, 12, 13, 23 Fagraea auriculata 24 
kyllingii 23 Ficus 15, 34 
radians 23 caulocarpa 6, 6, 7, 18 

Cystopteris 148 crassiramea 18 

Dahlia pinnata 169 deltoidea 18 

Davallia solida 24 drupacea 18 

Decaspermum paniculatum 18 fistulosa 19 

Dendrobium crumenatum 24 grossulariodes 19 
serra 24 hispida 15, 19 

Dentella repens 152 kurzii 6, 70, 11, 15, 19 

Derris scandens 4, 22 microcarpa 6, 9, 10, 11, 15, 19 
thyrsifolia 22 parietalis 22 
uliginosa 9, 22 stricta 6, 15, 19 

Desmodium umbellatum 18 sundaica var. beccariana 19 

Dianella ensifolia 23 superba 6, 7, 8, 9; 10:10, 1,45) 19pnae 

Digitaria ciliaris 23 tinctoria ssp. gibbosa 6, 6, 7, 19 

Diodia ocymifolia 152 variegata 4, 7, 19 
sarmentosa 152 Fimbristylis dichotoma 3, 25 

Diospyros 14 Flacourtia 15 
ferrea 6, 15, 18 Flagellaria indica 22 


Diospyros-Premna scrub 6 Furtadoa 53 


Index to Volume 38 


Furtadoa cont. 
mixtum comb. nov. 53 
sumatrensis 53 
Gaertneria acuminata 152 
grisea 152 
obesa 152 
vaginans 152 
viminea 153 
Gaillardia pulchella 170 
Garcinia hombroniana 19 
nigrolineata 14, 19 
angusta 153 
carinata 153 
florida 153 
griffithii 153 
jasminoides 153 
tubifera 153 
Gardeniopsis longifolia 153 
Gelonium glomerulatum 19 
Geophylla pilosa 153 
Gerbera jamisonii 170 
Glochidion littorale 19 
Gnetum gnemon 6, 15, 19 
latifolium var. funiculare 22 
Grewia paniculata 19 
Guettarda speciosa 19, 153 
Guioa pleuropteris 19 
Gymnacranthera 83 
bancana 201, 207 
paniculata var. zippeliana 83 
Gynochthodes coricea 22, 154 
sublanceolata 154 
Gynura procumbens 170 
sarmentosa 170 
Hedyotis auricularia 154 
biflora 154 
capitellata 154 
congesta 154 
corymbosa 154 
dichotoma 154 
diffusa 154 
herbacea 155 
pinifolia 155 
trinervia 155 
verticillata 23 
Helianthus angustifolium 170 
annuus 171 
tuberosus 171 
Hemiscolopia trimera 14, 15 
Heritiera littoralis 19 
Hernandia nymphaeifolia 19 
Heteroaridarum 43 
Hibiscus tiliaceus 19 
Homalomena 43 
gadutensis sp. nov. 45. 46, 48, 48 
gigantea 45 
egriffithii 50 
hastata sp. nov. 50. 5/ 
humilis 53 
megalophylla sp. nov. 43. 44, 45, 47, 48 
minutissima 53 
mixta 53 
padangensis sp. nov. 48, 49 
paucinervia 53 


237 


pendula group 48, 48 

rubescens 45 

rusdii sp. nov. 50, 52 

sagittifolia 47, 47, 48 
var. sumatrana 48 


Hopea-dominant forest 94 
Horsfieldia 69, 71 


affinis sp. nov. 210, 217, 219 
amklaal 80 
ampla 95, 97 
ampliflormis sp. nov. 95, 96 
amygdalina (Wall.) Warb. 188 
angularis sp. nov. 64, 65, 78, 97, 119 
angulata 110 
ardisiifolia 72, 73, 78 
aruana comb. nov. 100 
atjehensis sp. nov. 186 
australiana auct. 112 
australiana S.T. Blake 87, 88 
basifissa sp. nov. 64, 65, 98, 99, 109, 123 
borneensis 206 
brachiata var. sumatrana 75 
bracteosa 188, 195 
var. microcarya 191, 193, 225 
carnosa 198, 222, 223 
clavata sp. nov. 9/, 92, 95, 225 
corrugata 130, /3/, 134 
crassifolia 219, 220 
crux-melitensis 90, 91, 97, 225 
decalvata sp. nov. 125 
erubescens (in sched.) 112 
flocculosa 204, 207, 211,216, 219 
fulva 198, 199, 209, 222 
glabra (BI.) Warb. 64, 188, 198, 224 
glauca 188 
globularia var. minahasae 70 
gracilis sp. nov. 211 
grandis 203, 207, 208, 211, 213, 216, 219 
hellwigii 87, 121, 136, 138, 740, 142, 144 
hellwigii complex 137, 138 
key to the varieties 141 
hellwigii var. brachycarpa var. nov. 142 
var. hellwigii 121, 137, 138, 141, 740, 143, 
144 
var. hellwigii hybrid 141 
var. hellwigii X var. pulverulenta 137 
var. lignosa var. nov. 142 
var. novobritannica 121, 141 
var. pulverulenta 141 
inflexa sp. nov. 62, 63 
iriana sp. nov. 99, 101, 110, 119 
irya 55, 56, 78, 80, 102, 115, 121, 197, 198 
iryaghedhi 87 
laevigata 71, 97,99, 100, 101, 110, 112, 113, 
115, 118, 121, 122, 123, 126, 133, 134, 138, 
141 
key to varieties 117 
var. laevigata 117, 1/8, 120 
var. novobritannica comb. nov. 110, 117, 
118, 119, 120 
lancifolia sp, nov. 123, 1/24 
leptantha sp. nov. 137, 144 
macrocoma var. rufirachis 210 
macrocoma 75 


238 


Horsfieldia cont. 
macrothyrsa 188 
moluccana sp. nov. 65, 126 
var. moluccana 63, 66, 78 
var. petiolaris var. nov. 63, 67, 69 
var. pubescens var. nov. 68 
var. robusta var. nov 66, 68 
motleyi 206, 209, 211, 213 
novae-lauenburgiae 121 
novo-guineensis Warb. pro lectotype 100, 
101, 110 
novo-guineensis Warb., p.p. 110 
nunu 80 
obscurinervia 71 
olens sp. nov. 64, 80, 83 
olivaeformis 66 
pachycarpa 120, 130, 131, 133 
palauensis 78, 79 
pallidicaula sp. nov. 191, 225 
key to varieties 192 
var. macrocarya var. nov. 193 
var. microcarya comb. nov. 193 
var. pallidicaula 192, 193 
parviflora (Roxb.) Sinclair 65, 67, 69, 71, 72, 
74, 76, 78, 110, 115, 123, 125, 126 
paucinervis 212 
pilifera 101, 102, 110, 112, 118, 119, 121 
polyantha Warb. 99, 100, 101, 110 
praetermissa 132, 135 
psilantha sp, nov. 112, 115 
pulcherrima sp. nov. 202, 206 
pulverulenta 135, 137, 138, 144, 197 
punctatifolia 64 
ralunensis 87, 141, 143 
reticulata 213, 215, 216, 217, 218 
ridleyana 197 
rostrata 106 
roxburghii 67 
rufo-lanata 213, 216, 218 
sabulosa 185, /87 
salicifolia 87 
Samarensis sp. nov. 76 
schlechteri 95, 106, 107, 109 
sect. Orthanthera 87 
speikensis 64, 68, 81, 82 
ser. Punctatae 64 
sessilifolia sp. nov. 201 
sinclairii sp. nov. 90, 110 
smithi 74, 77, 80, 101, 102 
sparsa sp, nov. 188, 194 


spicata 59, 60, 66, 67, 74, 78, 80, 84, 101, 102,, 


115, 119, 120, 121 
var. sepikensis 64, 68, 69, 81 
var. spicata 68, 123, 132, 135 


splendida sp, nov. 202, 213, 217, 219, 213 2/4, 


217 
squamulosa sp. nov. 9/, 92, 93 
sterilis sp. nov. 191, 192, 197, 224 
subtilis 102, 196, 107, 109 
key to varieties 103 
var. aucta var. nov. 105, 106 
var. calcarea var. nov. 104 
var. rostrata 106 
var. schlechteri 95, 104, 109 
var. subtilis 103, 105, 106, 107 


Gard. Bull. Sing. 38 (1985) 


sucosa auct. 195 
sucosa (King) Warb. 188, 192, 195, 225 
key to varieties 189 
subsp. bifissa subsp. nov. 190 
var. microcarpa 191 
var. sucosa 189, 190 
superba 200, 202, 206, 207, 208 
sylvestris 84, 85, 87, 202 
var. villosa 87 
talaudensis sp. nov. 75, 77 
tomentosa 201, 213, 216, 219 
triandra sp. nov. 195, 196 
tristis sp. nov. 197 
tuberculata 66, 68, 113, 115, 126, 127, 130, 
129, 133, 134, 141 
key to varieties 128 
var. crassivalva var. nov. 130 
var. tuberculata 128, 133, 134 
wallichii 186, 204, 207 
warburgiana 74 
whitmorei 113, //6 
Hottarum 43 
Hoya diversifolia 11, 21 
Hydnophytum formicarium 24, 155 
Hypobathrum coniferum 155 
Hypodematium 148 
crenatum 148 
glabrious comb. nov. 148 
Ipomoea illustris 22 
pes-caprae 23 
Ischaemum muticum 23 
Isotoma longiflora 166 
Ixora chinensis 155 
coccinea 155 — 
congesta 19, 155 
finlaysoniana 155 
grandifolia 156 
javanica 156 
lobbii 156 
pendula 156 
Jackia ornata 156 
Kleinhovia hospita 19 
Knema globularia 6, 6, 9, 19 
laevigata 110 
malayana 20 
Lactuca indica 171 
sativa 171 
Lasianthus appressus 156 
attenuatus 156 
chryseus 156 
constricta 156 
cyanocarpus 23, 157 
densifolius 156 
ellipticus 157 
griffithii 157 
maingayi 157 
ridleyi 157 
scabridus 157 
singaporensis 157 
stipularis 157 
tomentosus 157 
Lastrea syrmatica 147 
Lastreopsis 148 
Laurentia longiflora 166 
Lecananthus erubescens 157 


Index to Volume 38 


Leea indica 20 
Lepisanthes fruticosa 20 
Litsea glutinosa 20 
Lobelia affinis 166 
zeylanica 166 
Loeseneriella pauciflora 22 
Lucinaea membranacea 158 
morinda 158 
Lumnitzera 3 
Lygodium 7, 9 
flexuosum 25 
Macaranga javanica 20 
Mallotus tiliaefolia 20 
Manikara 12 
kauki 6, 7, 15, 20 
Medinilla hasseltii 24 
Melampodium divaricatum 171 
Melastoma sanguineum 20 
Memecylon coerulum 6, 20 
ovatum 20 
myrsinoides 20 
Messerschmidia (Tournefortia) 12 
argentea 20 
Micropropagation of Lagerstroemia species: 
choice of nodal segments 177 
growth response to MS salt, IAA & Kinetin 
176, 177 
effect on shoots 178 
using nodal segments and shoot tips 180 
effect on rooting 
of IBA concentrations 177-179 
effect on shoot induction 
of BAP 117 
of BAP and 2ip treatments 176, 177, 178 
effect on rooting of excised shoots 
in agar medium with IBA & MS salt 178 
in sand medium with IBA & MS salt 179 
Mikania cordata 171 
micrantha 171 
scandens 171 
~ Morinda citrifolia 12, 13, 73, 20, 158 
ridleyi 158 
rigida 158 
umbellata 22, 158 
Mussaenda erythrophylla 158 
flava 158 
glabra 158 
luteola 158 
mutabilis 159 
philippica 159 
x Alicea, x Dona Aurora, x Dona Luz, 159 
Mussaendopsis beccariana 159 
Mycetia malayana 159 
Myristica aruana 100, 101 
guatteriifolia 6, 7, 16, 20 
laevigata 120 
nesophylla 101 
tomentosa 211 
Myrmecodia armata 159 
tuberosa 159 
Myrsine porteriana 20 
Myrtaceae-Vatica-Camnosperma forest 81 
Nauclea officinalis 159 
subdita 159 
Neolitsea zeylanica 20 


Nephrolepsis 6 
biserrata 11, 25 
Ochrosia 4 
oppositifolia 3, 16 
Oldenlandia biflora 154 
corymbosa 154 
dichotoma 154 
diffusa 154 
paniculata 3, 23, 154 
trinervia 155 
Oncosperma filamentosa 20 
Ophiorrhiza singaporensis 159 
Oxyceros fragrantissima 162 
longiflora 162 
penangianus 162 
scandens 162 
Paederia foetida 160 
scandens 160 
verticillata 160 
Pandanus 4 
dubius 5, 6, 6, 7,9, 12, 16, 20 
epiphyticus 16 
odoratissimus 20 
odoratissimus var. laevis 14 
spurius 14 
Panicum repens 23 
Paramignya umbellata 22 
Parinaria corymbosa 6, 6, 13, 13, 20, 41 
Parsonia spiralis 22 
Paspalum conjugatum 24 
Pavetta indica 160 
var. canescens 160 
Pedicellarum 43. 
Pemphis 3, 27 
acidula 3, 14, 20 
Pentaphragma x elliptica 167 
Pentaphragma horsfieldii 166 
ridleyi 167 
scortechinii 166 
Pentas carnea 160 
lanceolata 160 
Pertusadina eurhyncha 150 
Petunga conifera 155 
Phlebigonium impressum 147 
Phoebe declinata 20 
Phyllanthus? 16, 22 
Phyllanthus sp 6, 22 
Phymatarum 43 
Piper retrofractum 4 
Pisonia aculeata 22 
excelsa 16 
grandis 6, 9, /0, 11, 16, 20, 33 
grandis forest 36 
grandis wood 6, 8, 9 
Pithecellobium contortum 20 
ellipticum 20 
Pittosporum ferrugineum 21 
Planchonella firma 16, 21 
linggensis 16 
obovata 9, 21 
Plocoglottis porphyphylla 24 
Pluchea indica 171 
Podocarpus 4 
polystachyus 21 
Polyalthia sclerophylla 16, 21 


239 


240 


Polypodium scolopendria 13, 24 
Pometia 86 
Pometia-Intsia Forest 109 
Pongamia pinnata 21 
Porterandia anisophylla 161 
Premna corymbosa 16 
integrifolia 16 
obtusifolia 6, 11, 16, 21 
Prismatomeris malayana 160 
tetrandra 160 
Pseuderanthemum 7 
crenulatum 24 
Psidium 15 
Psychotria angulata 160 
cantleyi 160 
griffith 160 
helferiana 160 
maingayi 161 
malayana 161 
obovata 161 
ovoidea 161 
penangensis 161 
ridleyi 161 
rostrata 161 
sarmentosa 22, 161 
stipulacea t61 
Pteridium aquilinum 25 
Pteridris acutissima 147 
syrmatica 147 
Pteris ensiformis 25 
Randia anisophylla 161 
auriculata 162 
cochinchinensis 162 
exaltata 4 
fragrantissima 162 
longiflora 162 
macrantha 162 
macrophylla 162 
penangiana 162 
scandens 162 
schoemannii 4 
Rhizophora 4, 5 
conjugata 9 
Rothmannia macrophylla 162 
Rubiaceae, key to genera 149 
Rudbeckia serotina 172 
Sagenia 146 
Salacia chinensis 22 
Sambucus canadensis 166 
javanica 166 
Saprosma glomerulatum 163 
Sarcocephalus junghuhnii 159 
Sauropus albicans 22 
Sauropus sp. 22 
Scaevola 3, 6 
frutescens 167 
sericea 167 
taccada 21, 167 
Schefflera venulosa 11, 22 
Schizachyrium sanguineum 14, 24 
Scindapsus sp. 25 
Scleria lithosperma 24 
Scyphiphora hydrophyllacea 163 
Sigesbeckia orientalis 172 
Solidago altissima 172 


Gard. Bull. Sing. 38 (1985) 


Sparganophorus vaillantii 172 
Sphaeranthus africanus 172 
Spilanthes acmella 172 
Spinfex littoreus 14, 24 
Sporobolus virginicus 3, 24 
Stenochlaena 7, /3 
palustris 9, 13, 25, 36 
Sterculia foetida 6, 12, 21 
Stereospermum fimbriatum 21 
Streblus ilicifolius 4 
Stylidium tenellum 167 
Stylocaryna adpressa 163 
Styphelium 16 
Syndrella nodiflora 172 
Tacca leontopetaloides 5 
Taeniophyllum serrula 24 
Tagetes erecta 172 
patula 172 
Tarenna adpressa 163 
fragrans 163 
grandifolia 163 
lancifolia 163 
mollis 163 
ridleyi 163 
stellulata 163 
Tectaria 146 
key to species 147 
brachiata 146, 146, 147 
brachiata 146 
cherasica 146 
coadunata 145, 146, 147 
crenatum 148 
curtisil sp. nov. 145, 146 
fulcipes 146 
subtriphylla 147 
translucens sp. nov. 145 
variolosa 146, 146, 147 
Timonius compressicaulis 163 
finlaysonianus 163 
flavescens 164 
peduncularis 164 
wallichianus 164 
wrayi 164 
Tithonia diversifolia 173 
Tridax procumbens 173 
Ucaria attenuata 164 
cordata 164 
gambir 164 
glabrata 164 
jasminiflora 164 
longiflora 164 
ovalifolia 165 
pedicellata 164 
pteropoda 174 
roxburgiana 165 
sclerophylla 165 
Urophyllum glabrum 165 
griffithianum 165 
hirsutum 165 
macrophyllum 165 
streptopodium 165 
trifurcum 165 


VERNACULAR NAMES 
Abuino’o 129 


Index to Volume 38 


Vernacular Names cont. 
Aininiu, Ainynu 58, 129 
Airawikoepata 104 
Aitobi 104 

Akar subiak 170 
Ambuino’o, Ambuynor 129 
Amklaal 58 

Anunu magilioro 61 
Apaap 139 

Aragay 74 

Asem-asem 205 

Baa 134 

Babijag 98 

Baiwach 105 

Bale bale 129 

Bangera 83 

Bendoei 104 

Bepoes 98 

Berambong 150 
Betelohoi 98 
Beterohooi 67 
Boskomok 104 

Bunga china, Bunga susu 153 
Camarngur 139 

Cengan 163 

Cheem 108 

Cheeweng 124 

Dagoan 74 

Duguan 72 

Duria 225 

Euoe 110 

Fohja 139 

Gaben 122 

Gaigihab 111 

Gefrah 122 

Gosora 67 

Guma 97, 108, 122, 139 
Gumaga 136 

Hafringee 105 

Hamana 93, 111 

Hota 139 

Ibuumkwaraf 136 
linapo 104 

Ilis 110 

Isasir 81 

Jangkang paya 221 
Jesasir, Jisasir 81 

Kajoe darodong lomba 198 
Kajoe haroeja 221 
Kajoe penara 198 
Kamojer 67, 119 
Kamopi 64 

Kamore, Kamorei 68, 69 
Katumbi jantan 169 
Kawok-kawoe 119 
Kokotetepina 129 
Kolantie 70 

Krabo 172 

Kuleman, Kuleman 61, 67 
Kumpang balau 192, 218 
Kumpang ensuliue 221 
Kumpang sadara 221 
Kumpang tembaga 202 
Kumpang-perawan 186 


Kumpung 204 
Kupgne 134 


La gele kuku, Lagele kuku 121, 139 


Lagasi 74 
Laran’a 75 
Luhakon 119 
Madak 64 

Mag 104 
Mamasoh 139 
Mamgananieproi 122 
Mangaifa 104 
Ma-tak 81 
Mbowak 67 
Medak 69 
Medal 108 
Mengkudu 153 
Merambong 173 
Merampat 190 
Mong-mong 134 
Naufora 139 
Ngai camphor 168 
Niniwo 70 

Njet 104 
Numba 110 
Nungan 121 
Nunu 58 
O’hénga 139 
Oara 104 
Onguaka 61 
Oriomo 104 


Pala hutan, Pala utan 67, 78 


Patepa 136 

Peh (begie) 104 © 
Peita 119 
Penarahan 204 
Pérédah bésar 190 
Piangu, Pijangu 205 
Pive’ar 129 

Pohon lobi-lobi 78 
Poi 136 

Pokok beluntas 171 
Pokok german 170 
Posiposi 111 
Rengkéferék 104 
Rewwoh 104 
Roman 104 
Sabobo 136 
Saksak 111 
Samgoot 119 
Satim 205 


Sebohonggwa, Sebohongwa 67, 68 


Sekukumailor 126 
Selamae 81 

Sémies 69 

Serenai Laut 173 
Sodowa 132 

Simies 69 
Soemarallah oeding 205 
Soemarallah-falah 205 
Sumbong Jantan 170 
Sumbong 168 

Suri 104 

Ta’dara 221 

Tabenoe benoe 125 


241 


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Vernonia arborea 173 


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var. javanica 173 : - ) earn “it 
chinensis 173 -0t domea 


cineria 173 . . ott aiid aie 
patula 173 Pore 
Warszewiczia coccinea 165 
Wedelia biflora 173 
trilobata 173 
Woodsia 148 
Xanthium inequilaterum 173 
strumarium auct. 173 
Youngia japonica 174 
Zinnia elegans 174 
linearis 174 


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al AJAN 17 1986 ARNO 
LD ARBORETUM ISSN 0374-7859 
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x 
( THE GARDENS’ BULLETIN 
, SINGAPORE 
ee i ee 
X VOL. 38 (Part 1) Ist June 1985 
e CONTENTS 
PAGES N 
30) ey TSS Sap ES: eee 
The Botany of Some Islets East of Pahang and Johore .............:2.--+ssseeeeeeereeeenneeetess 1-42 
HOTTA, Mitsuru: 
New Species of the Genus Homalomena (Araceae) from Sumatra with a Short Note on 43-54 
the Genus Furtadoa 


eee veka nea an en ia he sae be hee see ee pee ee oe OP PRN AEE eee ae Pas 2 Ch RRR Peco ses oe we 


WILDE, W. J.J. O. DE: 


A New Account of the Genus Horsfieldia (Myristicaceae), Pt 2 ............-seseeeeeee eee eeeee 55-144 Y, 


Published by the Botanic Gardens 

Parks and Recreation Department 

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THE GARDENS’ BULLETIN 


VOL. 38 (Part 1) Ist June 1985 
CONTENTS 
PAGES 
CORNER, E. J. H.: 
The Botany of Some Islets East of Pahang and Johore .................0. cece eee eee 1-42 
HOTTA, Mitsuru: 
New Species of the Genus Homalomena (Araceae) from Sumatra with a Short Note on 43-54 
ie Genus uriadogs. 2:6. 62.0.8 2251.8. We AS RESET at LIK Pat Khare EP nL ak fda Sak ago 
WAIRDE AW is JcOrpe: 
A New Account of the Genus Horsfieldia (Myristicaceae), Pt 2 ..............0.. cece eee ee eee. 55-144 


Published by the Botanic Gardens 

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Ministry of National Development 
Cluny Road, Singapore 1025. 


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The Botany of Some Islets East of Pahang and Johore 


Ba fi. Comer 
91 Hinton Way, Great Shelford 
Cambridge CB2 5AH, England 


Abstract 


The appearance and vegetation of the islets Babi Tengah, Baru, Berhala, Chibeh, Duchong, Gup, 
Labas, Pyah, Rengis, Sepoi, Setindan, Tokong Burong, and Tulai are compared with Tanjong Ruit in 
Tiuman, as they were in 1932-1936. Their status as degenerate headlands in the Riouw Pocket is 
considered. The islets belong to three geological formations. Remains of the giant clam, Tridacna, were 
found on Pulau Tulai. 


This is a historical account of the vegetation of the small islands to the west and 
south of Pulau Tiuman (Tioman) and off the Mersing coast, as I found them fifty 
years ago. In June 1932 and August 1935 I accompanied Mr William Birtwistle, 
Officer-in-charge of Fisheries F.M.S., on his inspection of fishing boats and fishing 
villages on the Pahang coast. We visited most of the islets on one or other day from 
early morning till 1 p.m. The ocean swell would then rise and squalls descend, and 
we would have to return in the rowing boat to the launch, either the Sri Gala in 
1932 or the Sri Pekan in 1935; generally, the shore was too rocky for safe approach 
except in calm sea. The first trip was exploratory, for no one could tell me what the 
islands looked like or what vegetation they had; the second was more thorough. 
There was usually time to investigate fully in the one morning the vegetation of one 
islet. Collections were made of plants that I could not identify for certain and they 
have been deposited in the Singapore herbarium. Uncertainty of names has de- 
layed publication but, now, with many revisions in the Flora Malesiana, edited by 
Professor C.G.G.J. van Steenis, most problems have been resolved. I have fol- 
lowed that botanical nomenclature and have, accordingly, omitted the burden of 
citation of authors. 


On the second trip I realised that most of these islands corresponded with granite 
headlands of Tiuman in various degrees of disintegration. I have used, therefore, 
the headland of Tanjong (Tanjung) Ruit at the north end of Juara Bay, on the east 
side of Tiuman, as the standard for comparison. 


At the end of this article I have given notes, mainly geographical, on some of the 
more characteristic plants. The whole region is the northern part of the Riouw 
(Riau) Pocket (Corner 1978); the plants are vestiges of its history. If their ex- 
traneous distribution is plotted, not as circular or elliptical areas but as outline 
tracks, more in the manner of Croizat’s Panbiogeography, it will be seen that they 
enter the region from all points of the compass, as if they were comets orbiting 
about it. 


Tanjong Ruit, Juara Bay, Pulau Tiuman 
Figure 1, plates 1 & 2 


Tanjong Ruit ends in a low rocky point, about 30 ft high, of granite boulders. It is 
connected with the main island by a neck or isthmus, about 50 yards long, com- 


Editorial Note. Current spellings of place names, if different, are inserted in parentheses following the 
old spelling on first mention in the main text. 


Be 
: 
= die oe 
3 
Pp 
R = 19 Ta 


Fig. 1. Map of the islands about Pulau Tiuman: scale, 1.5 in = 10 miles. B, P. Berhala; Ba, Baru Rock; 
Be, Bebar; C, P. Chibeh; D, P. Duchong; E, Endau; G, P. Gup; J, Juara Bay; L, P. Labas; P, 
P. Pyah; R, Rompin; S, P. Sepoi; Sr, P. Sribuat; T, P. Tulai; TB, P. Tokong Burong. 


posed of small water-worn granite boulders just above high tide level, but probably 
awash at the highest tides and in storms. The island here descends into a steep 
granite face plunging into the sea. It leads westwards for about a third of a mile to 
meet the sandy bay where it turns, as it were, inland. I explored the forest that 
bordered on to this steep face, the isthmus, and the rocky point. 


The bare granite face varied 12-35 ft high between the high tide level and the 
edge of the forest. It consisted of steeply sloping blocks of granite, broken up in 
places into irregular ledges and, where the face was breaking up, there were big 
boulders piled precariously on one another. At the foot of the face there was a talus 
of water-worn boulders exposed at low tide; a few big isolated boulders stood as 
rocks in the sea. Above the face the forest stretched inland on the steep hillside, 
with a moderately thick layer of earth and humus. The granite face, itself, swept by 
waves during the north-east monsoon, was bereft of vegetation except for some 
ledges and rock hollows in its upper part, and they supported small grasses, sedges, 
and other herbs. This bare granite face was a feature of most of the granite islets 
which plunged into the sea, its depth ranging from 17 fathoms off P. Sepoi to 20 
fathoms off Tg Ruit. I limited my exploration to the Terminalia-zone which was the 
front of the forest at the top of the granite face, and to the zone of Eugenia grandis 
immediately behind. There was no time to explore further inland. 


The Terminalia-zone was a broken and almost single file of trees and climbers 
with some shrubs and herbs, typical of the Terminalia-Barringtonia formation on 
the rocky east coast of Malaya. Here and there, in sheltered places, there were 
intrusions of the inland forest. The seeds and fruits of many of these plants are 
water-borne and, presumably, are splashed up the granite face by waves. The list of 
plants in this zone is given in Table 1. It will be seen that there were no Cycas, 


2 


Botany of eastern Pahang and Johore islets 3 


Lumnitzera, Peltophorum, Scaevola, or Casuarina, which belong to the sandy or 
less boisterous coasts. 


A striking feature of this Terminalia-zone, as on other rocky parts of the east 
coast, was the great abundance, often thick masses, of such epiphytes as orchids, 
aroids, and ferns growing on the rocks. Conditions of insolation, drainage, and 
intermittent rain seem comparable with those of the limbs of trees, and the epiphytes 
which can stand the sea-spray become truly saxicolous. Climbers, also, where they 
could root into a crevice between the exposed boulders, spread thickly over them in 
a mat or carpet that hid the boulders, curtained the sides, and could be lifted off. 
Similarly, several species of strangling fig, normally beginning as epiphytes, car- 
peted the exposed rocks without developing erect branches; such carpets were 
mostly sterile and it was not till many years later that I managed to identify the 
species more or less satisfactorily. 


Another feature of the granite face was a special habitat for small herbs where 
some humus and rain-water could collect in crevices and small hollows, especially 
where shaded by trees. Some of the hollows were merely little basins in the rock 
from which the plants could be scooped, as if from a pot. Such plants were 
Aneilema (SFN 29794), Cyperus cyperinus, C. dubius, C. javanicus, Fimbristylis 
dichotoma, Oldenlandia paniculata, Sporobolus virginicus, and Vandellia hirsuta. 
This habitat was noticeably absent from the granite islets around Tiuman; the 
granite face might have been too wave-swept or have been unable to accumulate 
enough soil or rain. . 


Immediately behind the Terminalia-zone there was that of Eugenia grandis, 
mostly 1-2 trees deep. The trees did not actually front the coastal forest but 
occurred where there was shade enough for their roots. The seeds of E. grandis 
germinate in the open or in shade but seem unable to establish themselves in high 
forest or in the shade of their own kind. Thus, the trees start in the open and come 
slowly to overshadow the Terminalia-zone. However, this coastal fringe of forest 
was retreating, as witnessed by the fallen boulders, and the interaction between E. 
grandis and the Terminalia-zone was by no means clear. The inland forest intruded 
with seedlings and saplings into the zone of E. grandis. The coastal forest at Tg Ruit 
was thinned to a strip and, on the islets around, it was variously disrupted as they 
were disintegrated. The headland had two special features. 


The point itself carried the Terminalia-Barringtonia formation which stopped 
abruptly at the isthmus of water-worn boulders. These were mixed with coral debris 
and sandy detritus, heaped together by the waves as a low barrier. It carried an 
almost pure stand of Pemphis acidula, which did not occur elsewhere at the 
headland. It formed twiggy gnarled bushes, up to 12 ft high, the leaning and twisted 
trunks rooting in the debris. This sort of junction of a headland with a spit of talus 
seems to be a favourite habitat of Pemphis. On the rocky mainland coast of east 
Johore, with haematite quartzite shale, Pemphis forms prostrate sprawling mats, 
scarcely a foot high, and rooted in the crevices, but I did not see it in this manner on 
the granite boulders of the Tiuman complex. 


Then at the landward end of the isthmus, where the granite face rose steeply. 
there was a little wood of Ochrosia oppositifolia, about 50 yards long and 30 yards 
wide, between the Terminalia-zone and that of E. grandis. The trees, up to 50 ft 


tall, were thickly placed, rather slender, and together with many of their seedlings 
and saplings formed an almost pure stand. The ground was a mixture of small 
boulders and sand at the foot of the cliff. It was difficult to see why such a stand 
should have developed. The species did not occur in the Terminalia-zone on the 
granite face; on the east coast of Johore it frequented sandy shores. It seemed to be 
the result of a freak current or storm which had washed up the fruits on this part of 
the beach at a time when it was bare of vegetation, and that the colony had 
persisted while the Terminalia-zone took over the frontage. 


Pulau Tulai 
Figure 1 

This is the largest of the small islands off Tiuman and lies about 3 miles WNW. of 
its north end. In 1935 it was largely covered with inland forest. I was able to explore 
in detail only the NW. coast and the bay on the west of the island. The whole coast, 
except for this bay, is rocky and composed of granite blocks c. 20 ft high in situ and 
others tumbled on top of them, and on the NW. side, at least, there was the talus of 
water-worn boulders plunging into the sea without foreshore. The coastal flora was 
essentially the same as at Tg Ruit but without Ochrosia; there was the fringing 
Terminalia-zone along the top of the granite face with an interrupted zone of E. 
grandis connecting with the inland forest. There were, however, the following 
additions:— 


Adenanthera pavonina (strangely none seen at Tg Ruit), 

Cissampelos sp., 

Clerodendron inerme, 

Colubrina asiatica, 

Derris scandens, 

Elatostema sp. (on rocks in the shade), 

Ficus variegata (in E. grandis zone), 

Hemiscolopia trimera (common on rocks in the Terminalia-zone), 

Piper retrofractum (on shaded rocks with Elatostema, often in thick carpets 
sweeping up vertical sides of rocks), 

Randia schoemannii (R. exaltata), 

Streblus ilicifolius (not on exposed rocks), 

Xylocarpus (Carapa) moluccensis (a frequent spreading tree). 


Hydnocarpus ilicifolia was abundant as a small tree to 30 ft high both in the 
coastal fringe and in the inland forest. There were no Cycas, Pandanus, or Podo- 
carpus. 


The main bay, as a bight on the west side of P. Tulai, had a mangrove forest of 
considerable extent. Here the granite face receded inland and, at the SE. corner of 
the bay, a small sluggish stream flowed over a shallow flat of coral detritus, exposed 
at low water, before emptying into the bay. The shallow flat was roughly semicircu- 
lar and about 200 yards in diameter. Here was the mangrove forest composed of 
Rhizophora conjugata, R. mucronata, and Bruguiera gymnorrhiza, on clean firm 
ground without the usual mud and slime. There were many coral fragments mixed 
with the sand and in places lumps of dead coral more or less buried in situ. R. 
mucronata was by far the commonest, occupying most of the flat and all the 
seaward front, but scarcer inland. R. conjugata occurred sparingly on the landward 
side. B. gymnorrhiza was scattered in the central part of the flat and abundant on 
its landward side, even in places to the exclusion of Rhizophora. Some trees of B. 


Botany of eastern Pahang and Johore islets 5 


gymnorrhiza stood 60-70 ft high, those of R. mucronata being somewhat smaller. 
Pneumatophores were present in great abundance and, at low water, it could be 
seen that a tangle of Rhizophora roots flanked the stream; at high water a prahu 
could float down it. The mangrove was advancing into the bay which was sheltered 
from the NE. monsoon and from the SE. tenggara*. How old this mangrove was, I 
could not determine, but it was growing on a fringing reef that must have been 
raised in fairly recent times. 


Immediately behind the mangrove flat, there was a sandbank 4-5 ft high, 
stretching inland, and evidently a former beach. It was largely planted with coco- 
nuts but had clearly been covered originally with the Terminalia-Barringtonia 
formation and that of E. grandis; there were many plants of Tacca leontopetaloides. 
The sandbank led to the granite slope with inland forest. 


Just north of the mangrove flat, by some large rocks at high tide level, there were 
several large shells of the giant clam Tridacna, 2-3 ft across, in situ and upright but 
almost completely buried in sand; the wavy outline of the valves was more or less 
visible. Out in the bay we saw through its clear water smaller living clams about a 
foot wide at depths of 8-10 ft; we also met them in our early diving efforts with 
helmet and air-pump in this bay. They lived close to the rocks, neither in the sand 
of the bay nor in the living coral. Here was further evidence that P. Tulai had been 
raised or tilted, perhaps some 20-30 ft, to uplift and expose the giant clams. 
Previously there could have been no mangrove there, and where the stream flowed 
there might have been a narrow strait separating the SW. part of the island from the 
rest. As this part was raised, so the bay silted up to give the sandy padang with 
coconuts. In other words, the age of the mangrove must be connected with the 
raising or tilting of the island. 


No one lived on P. Tulai but it was frequented by fishermen and, doubtless, the 
owner of the coconuts. There was a well, supplying fresh water, on the south side of 


the mangrove. From the general appearance of the forest, it seemed that the island 
had never been inhabited. 


The broad-leafed and tree-like Pandanus dubius was abundant on the west side 
of P. Tulai. It grew on the sandy shore of the bay in the Terminalia-zone and in the 
hillside forest somewhat inland, but not on the rocky coast. The biggest plants had 
stems up to 25 ft high and were not so large as those on P. Chibeh (Cibeh). I saw it 
at the north end of Tiuman and on the south side of Juara Bay, but not at Tg Ruit, 
and I did not see it on the south and west of Tiuman. | found no flower or fruit 
anywhere in August 1935, but its seedlings were abundant. 


Pulau Chibeh 
Figures 1 & 2, plates 3-6 


This small island, about 250 ft high, lies a mile or so north of P. Tulai. It consists 
of immense granite blocks more or less in situ, immense fallen boulders, a bare 
granite face above tide level, and more or less of a submerged talus of boulders and 
pebbles. It resembled a disintegrated headland of Tiuman and its vegetation was 
that of Tg Ruit on a diminished scale, but without regular formation (Table 1). I 
visited it on 19 August 1935 from 7 a.m. till 1 p.m. when the swell forced me to 


*Tenggara”’ means south-east but as used by the fishermen of the East Coast, the use is transferred 
from the direction to the afternoon wind itself on which they sailed home. Ed. 


6 Gard. Bull. Sing. 38(1)(1985) 


return to the launch. I explored most of the island but did not cover the whole of 
the exposed east side where the immense boulders made passage very difficult. 
However, they were largely bare of vegetation and I did not miss any plant of 
importance. 


The north and south ends of the island were steep with the boulders exposed and 
hot, but on some of them there were mats of saxicolous strangling figs with tufts of 
~ Cyperus javanicus and Nephrolepis in crevices, as the only vegetation. The speci- 
mens which I collected resolved into F. kurzii, F. stricta, and F. tinctoria ssp. 
gibbosa, not F. microcarpa as I had supposed; all occur, together with F. microcar- 
pa, on rocky parts of the east coast of Malaya and they show no obvious ecological 
preferences. Higher up on the north end of the island there were scrubby patches of 
Diospyros ferrea, Memecylon coeruleum, and Premna obtusifolia. Then, in shel- 
tered nooks on the north and west sides, near to the sea-level, there were a few 
bushes of Scaevola. 


Fig. 2. Sketch map of the main vegetation on P. Chibeh (left) and P. Sepoi (right). A, Atalantia wood; 
EP, Eugenia grandis and Pandanus dubius; Fc, Ficus caulocarpa; Fs, Ficus superba; Ft, Ficus 
tinctoria ssp. gibbosa; f, Diospyros-Premna scrub; Ma, Manilkara kauki; My, Myristica guatter- 
iifolia; Pa, wood of Parinari corymbosa; Pi, Pisonia grandis wood; R, Calamus chibehensis; Y. 
Phyllanthus sp., SFN 29848; +, Knema globularia. 


On the east side, above the bare granite face, there was a rather open wood of 
Eugenia grandis with other trees of the Terminalia-zone and abundant Pandanus 
dubius. On the west side, which was sheltered and not so steep, this kind of 
woodland with trees up to 50 ft high was better developed and had a frontage of 
Pisonia grandis, the white branches of which were conspicuous from out at sea. The 
woodland extended over the centre of the island and consisted mainly of tall trees 
of E. grandis, Parinari corymbosa, Sterculia foetida, Vitex pubescens, and Ficus 
caulocarpa with F. stricta; the smaller trees were Atalantia monophylla, Caryota 
mitis, Eurycoma longifolia, Gnetum gnemon, Hydnocarpus ilicifolia, Knema globu- 


Botany of eastern Pahang and Johore islets 7 


laria, and Pandanus dubius (up to 40 ft high with stilt-roots up to 12 ft high). 
Surprisingly, the herb Pseuderanthemum was very abundant in this woodland. 
There were three plants of special interest in the upper part of the island. At the 
north end there were two trees of Manilkara kauki, up to 30 ft high, standing apart 
from the woodland. In the woodland in the centre of the island there were one or 
two trees of Myristica guatteriifolia. Then, in the north-west corner of the woodland 
and in the south part, there were two clumps of the rotan which C.X. Furtado 
called Calamus chibehensis. Beside the northern clump of rotan there stood two 
big trees of Ficus tinctoria ssp. gibbosa and one of F. caulocarpa which had a nest of 
the sea eagle. Another big tree of F. caulocarpa stood on the outside of the 
southern clump of rotan. I noted two trees of Ficus variegata in the woodland. It 
was an extraordinary association of forest relics. 


The trunks of the strangling fig-trees stood directly on the boulders over which 
their roots spread and entered into crevices, possibly down to the water-table (? 
brackish) in the centre of the island. The roots were so numerous and strong that 
they undoubtedly helped to basket the loosening boulders and hold them up. How 
the fig-trees had started was not evident. If they had begun as epiphytes, which 
their trunks suggested, there was no trace of the host-tree. I did not meet Man- 
ilkara anywhere else on the islets off Tiuman; it was not likely to have been 
overlooked because of its appearance like a chiku tree with dark fissured bark and 
white undersides to the stiff shiny leaves. At the time of my visit in August, the 
trees of F. caulocarpa and Pisonia were getting new leaves. 


The climbers and ferns mostly formed carpets on the boulders, both in the open 
and in the lighter shade of the woodland. Their variety added to the peculiar 
composition of the flora. 


I concluded that P. Chibeh had a woodland made largely of plants of the E. 
grandis forest of rocky coasts, with a few relics of inland forest, as shown by 
Manilkara, Myristica, and F. variegata. 


I noted that ants were very abundant, especially the red keringga with fiery bite. 
There were several large ant-hills or mounds up to 3 ft high. 


Pulau Sepoi 
Figures 1-4, plates 7-13 


This is a single rounded hill-top, 230 ft high, three miles due west of P. Tulai, and 
the sea-depth around is given as 17 fathoms. It is a mass of granite blocks and 
boulders similar to P. Chibeh and rather smaller. Its vegetation was also similar but 
with still fewer species (Table 1). I visited it in June 1932 and on 20 August 1935. It 
was tenanted by many terns which nested on the higher rocks where the vegetation 
began; they laid their eggs, apparently one to a nest, on bare ground under a 
projection of the rock without any sticks or leaves. There were numerous rocky 
caves round the foot of the island, as on P. Pyah (Paya), but which I did not note on 
P. Chibeh. The upper part of the island was covered with a low wood, 30-40 ft high, 
from which some large fig-trees projected, notably Ficus superba. The top con- 
sisted of large and small boulders with very little, if any, soil. The trees were well 
spaced, mostly on the boulders, with little or no undergrowth so that it was easy to 
walk among them. The granite face between the sea and the vegetation had 
scattered tufts of Cyperus javanicus and some sprawling patches of Lygodium and 
Stenochlaena. 


\ : 


Fig. 3. Sketch of a transect from west (left) to east across P. Sepoi. A, Atalantia wood; C, mats of 
climbers on the rocks; Fs, Ficus superba; Pi, Pisonia grandis wood; V, Vitex pubescens. 


i 


Fig. 4. Sketch of a transect from south (left) to north across P. Sepoi. Letters as in Fig. 3; Ac, 
Allophylus cobbe. 


Botany of eastern Pahang and Johore islets 9 


The woodland had two very different parts. That covering the upper part of the 
island I called the Atalantia wood because of the abundance of A. monophylla. The 
other was the Pisonia wood on the west and south-east parts of the island, on the 
lower slopes where there was some shelter from the NE. monsoon and the slopes 
were less steep. The Atalantia wood had also abundant Hydnocarpus ilicifolia, 
Planchonella obovata, and Ficus superba, the many roots of which clasped the 
rocks and descended into the depths of the granite. The fluted trunks of A. 
monophylla, set with thorny twigs, were easily recognisable. Near the top, in the 
Atalantia wood, there were numerous trees of Knema globularia. At its outskirts in 
the south-west, Derris uliginosa formed close mats on the rocks. Several large trees 
of Vitex pubescens, up to 50 ft high, occurred on the east side of the wood where 
there was the one patch of the small climber Phyllanthus? (SFN 29848). In contrast, 
the Pisonia woods were almost pure stands of P. grandis with an undergrowth of 
Stenochlaena palustris and a few trees of Adenanthera pavonina, Ficus microcarpa 
(with many aerial roots from the branches), and some stray Atalantia. The smooth 
grey trunks of Pisonia were seated on the rocks, with roots spreading over them in 
the manner of fig-roots. All the Pisonia trees were, in August, getting new leaves 
and beginning to flower, as on P. Chibeh; their white branches rendered them 
conspicuous. The trees of F. superba were also getting new leaves. 


In many places there appeared to have been landslips or, at least, the fall of 
boulders which had destroyed parts of the Atalantia wood, and masses of creepers 
were covering the fallen rocks. Remarkably, there were no Eugenia grandis, palms 
of any kind, Pandanus dubius, Terminalia catappa, Apocynaceous climber Aganos- 
ma, or epiphytes. In fact, there was no vestige of either the formation of E. 
grandis or of Terminalia. 


In August 1935, many of the trees showed signs of wilting. What little soil there 
was between the boulders was dry; fallen leaves crackled under foot. Patches of 
Lygodium and Stenochlaena in the open were brittle. The leaves of many smaller 
trees and their saplings were drooping, wilting or dried out. At first, I thought that 
this might be due to a landslip upsetting and breaking roots, but there was no actual 
indication of a recent fall. Then, I thought that the wilting might have been caused 
by the abundance of guano round the nests of the terns, but I could discover no 
relation between the nests and the wilting trees. The only cause seemed to be, 
simply, the lack of rain. The roots of the fig-trees, of Pisonia, and of Vitex 
pubescens penetrated deeply into the mass of boulders and their leaves showed no 
signs of wilting. 


There were many old and rotting trees, standing and fallen, in both the Atalantia 
and the Pisonia woods, where there were also plenty of seedlings and saplings. 
Hence I concluded that the vegetation of the island must have been of long 
standing. Landslips, boulder-falls, and periods of excessive drought evidently killed 
some of the trees and enabled a new generation to arise. Pigeons and fruit-bats, 
even the large hornbills, visiting the island from Tiuman or P. Tulai, might carry 
seeds to re-stock, but all the evidence that I could gather pointed to the conclusion 
that P. Sepoi was a degenerating headland of ancient coastal forest. Indeed, this 
cluster of islets about P. Tulai probably made long ago one hill or mountain which, 
in its turn, may have been part of Tiuman itself. I was thinking of subaerial 
denudation. Lowering of the sea-level during glaciations would have exposed the 
base of the hills without necessarily a connection. 


10 Gard. Bull. Sing. 38(1) (1985) 


Pulau Labas 
Figure | 


This is the fourth and smallest island of the P. Tulai group. It lies about two miles 
SW. of P. Tulai and about one mile SE. of P. Sepoi. I visited it also on 20 August 
1935. It is a collection of large and enormous granite boulders raised about 30 ft 
above the sea-level. There were scattered trees of Ficus superba, mats of F. 
microcarpa and Aganosma marginata on some rocks, and scattered clumps of 
Cyperus javanicus and the mangrove fern Acrostichum aureum. Otherwise, the 
only other ‘higher plant’ on the island was a tall fruiting coconut palm, evidently 
planted, with several seedlings around it. The island seemed to represent the stump 
of a headland to which, in time, P. Sepoi would be reduced. However, the presence 
of Acrostichum brought in another factor to suggest that P. Labas might be the 
remains of a small granite headland, such as at Tg Ruit, beside a mangrove forest 
such as on P. Tulai. Indeed, this mangrove at P. Tulai was probably the vestige of a 
much more extensive mangrove forest when all four islands were joined in periods 
of glaciation. 


All four islands were, and should be allowed to remain, historical sanctuaries. 


: me 
~as—0 a ye te 


Fig. 5. Sketch of a transect of P. Rengis. Ac, Allophylus cobbe; C, climbers on the rocks; Fk, Ficus 
kurzii; Fs, Ficus superba; Pi, Pisonia grandis; V, Vitex pubescens. 


Pulau Rengis and P. Pyah 
Figure | 


The very small island P. Rengis (Renjis), about 80 yards, across lies about a third 
of a mile from the west coast of Tiuman in Telok (Teluk) Tasek. I visited it on 16 
August 1935. It was a mass of granite boulders piled on top of one another as P. 
Labas, and with water-worn boulders strewn at the base. The island was tenanted 
by frigate birds, not nearly so numerous as the terns on other islets, and the guano 
was not so persistent; nevertheless, many trees were white with guano. The flora 


Botany of eastern Pahang and Johore islets 11 


was very limited to 11 species. There was no Terminalia-zone or Eugenia grandis, 
and I saw no herbs, grass, or sedge. It consisted of:— 


Allophylus cobbe var. marinus, common by the shore, 
Callicarpa longifolia, as a few scattered bushes, 

Ficus kurzii, as a few small spreading trees at the south end, with copious aerial 
roots, starting as a bush on the rocks; (easily mistaken for F. microcarpa), 
Ficus superba, as frequent big trees in the centre of the island, never as a 

prostrate mat, 
Hoya diversifolia, very common on trees and rocks, 
Nephrolepis biserrata, a single clump, 
Pisonia grandis, as the commonest tree by the shore, 
Premna obtusifolia, as a scraggy subscandent bush, 
Schefflera venulosa, abundant on trees and rocks, all its leaves white with guano, 
Trema amboinensis, as a few scraggy treelets, 
Vitex pubescens, as a single tree SO ft high. 


On 11 June 1932, when we had anchored for the night in Ayer (Air) Batang bay, 
I visited the islet P. Pyah, just off the south headland of the bay, but I failed to 
make an inventory of its flora. It was, as P. Rengis, a cluster of granite boulders on 
which the chief tree was Ficus superba. There were many caves round the foot 
where I spent most time looking for algae, but with little success. 


Pulau Tokong Burong, Baru Rock, and P. Gup 
Figure 1, plates 14 & 15 


A cluster of three steep islets of haematite shale or laterite-looking rock, about 
10 miles SW. of P. Sepoi and some 17 miles from Tg Penyabong on the mainland, 
make the group of Tokong (Tukong) Burong. I visited them on 16 August 1935. 
The largest rises to 160 ft high and the base, strewn with large boulders, reminded 
me of Tg Sedili. There was no forest. The islands were tenanted by thousands of 
terns which nested under tussocks of the grass Chrysopogon fulvus, as the only 
vascular plant on the islands. It grew from high tide level on the south face on 
terraces or niches in the rock, and right over the summit. There was no sward 
because the runways of the terns kept the tussocks more or less apart. There were 
no seedlings of other plants, stumps, or traces of others, from which it seemed that 
the islands, as they now stand, might never have been forested. The top of the large 
island had the appearance of having been burnt but, if so, the effect had merely 
halted the grass temporarily. 


Baru Rock, or P. Baru, and P. Gup are islets of roughly the same size, compara- 
ble with the largest of the Tokong Burong group. Baru Rock, 190 ft high, lies about 
11 miles south of Tokong Burong. P. Gup, 140 ft high, lies to the east in the same 
latitude and about 4 miles south of Tiuman. I was never able to visit either of them 
but, on a voyage to Kemaman in October 1935 in S.S. Mahidol of the Danish East 
Asiatic Company, I persuaded the captain to pass as close as possible to the islands 
so that I could have a look at them. He enjoyed the manoeuvre and handed me his 
telescope. The difference between the islands was as astounding as that between 
Tokong Burong and Sepoi. Baru Rock was a firm consolidated mass of rock, 
probably the haematite shale, with precipitous sides, serving as a ternery, and rising 
straight from the sea with few or no boulders at the foot. The upper two-thirds were 
covered with a tussock grass, presumably the same as on Tokong Burong. At the 


12 Gard. Bull. Sing. 38(1) (1985) 


north end, half-way up the island, there were a few small stunted trees with rather 
large leaves, which could have been Morinda citrifolia. Thus Baru Rock may have 
been less degraded than Tokong Burong. Perhaps both were originally forest-clad 
like P. Duchong (Ducong). 


In complete contrast, P. Gup was a mass of granite boulders with the upper part 
rather closely wooded, very similar to P. Sepoi. I could not distinguish any floristic 
details but looked unsuccessfully for the white branches of Pisonia. 


Pulau Duchong 
Figure 1, plate 16 


Two small islands lie near the mainland just south of Pontian. The larger, P. 
Duchong Darat, 80 ft high, is about a mile off the coast. The smaller, P. Duchong 
Laut, is about half a mile SE. of the larger. I visited Duchong Darat on 22 August 
1935, but had no time to visit the smaller, which did not appear to differ floristical- 
ly. They are steep craggy masses of haematite shale, similar to P. Setindan off 
Mersing and to Tg Sedili. The flora was that of the Terminalia-Barringtonia zone 
and of Eugenia grandis forest with some additions as relics, perhaps, of inland 
forest. Thus, in a general way, it resembled the coastal flora of Tiuman but several 
conspicuous plants were missing, namely the trees Atalantia, the strangling figs, 
Hydnocarpus ilicifolia, Manilkara, Pandanus dubius, Pisonia, and Sterculia foetida, 
the climber Aganosma, and the sedge Cyperus javanicus. The floristic list is given in 
Fable 1. 


Round the foot of the island there was the Terminalia-zone. The rest was covered 
with rocky forest with trees up to 60 ft high, though those of the upper part were 
merely 15-20 ft. Exposed rocks were covered with masses of sprawling climbers and 
epiphytes among which Asplenium nidus-avis was conspicuous. There was no 
evidence that the forest had been cut over. Various herbs and grasses of the sandy 
shore of the mainland occurred in small sandy places among the lower rocks. The 
Chrysopogon of Tokong Burong did not occur. 


Pulau Setindan 


I visited this island off Mersing on 15 August 1935 and on 30 January 1937. 
Geologically and floristically it was similar to the headlands of the east coast of 
Johore and to P. Duchong. There were the formations of Terminalia-Barringtonia 
and of Eugenia grandis, but where there had been inland forest, I found mostly 
scrub; at some time the island had been extensively cut over. The plants which I 
found are given in Table 1, for comparison with P. Duchong. The islands were 
generally similar but Setindan is larger and more representative; yet, there were 
unaccountable differencees. Certainly, Setindan relates more with the flora of the 
Riouw Pocket, as shown by the abundance of the southerly Tristania obovata. The 
lists in Table 1 largely repeat what I have written about the Sedili coast, but I 
decided to include it as a record what deforestation is likely to obliterate. It 
reminds one of what might be expected. 


Botany of eastern Pahang and Johore islets 13 


Pulau Berhala 
Figures 1 & 6, plates 17-21 


This islet lies north-west of Tiuman and about 20 miles north-east from Bebar on 
the Pahang coast. It is a flat granite platform, roughly circular, about 400 ft wide, 
raised a foot or more above sea-level and, probably, awash at highest tides or in 
storms. The platform is dissected radially by deep gullies plunging into the sea 
(12-13 fathoms deep). In the centre there was a sandstone mass, about 80 ft high, 
shaped like the crown of a hat to which the granite platform was the brim. It was 
neither the mass of granite boulders of the Tiuman islets nor the compact haematite 
of the coastal islets. The sandstone was fairly hard and distinctly, though not 
deeply, undercut round the granite platform. The islet clearly belonged with 
another geological formation and recalled Gunong Panti in east Johore. 


The vegetation was confined to the upper half of the sandstone hillock where it 
was rather evenly dense, but the granite platform was destitute of any plants 
except, perhaps, some lichens and microscopic algae. The flora consisted of merely 
5 species, thus: 


Stenochlaena palustris and Polypodium scolopendria in a low tangle, with tufts of 
Cyperus javanicus, forming the lowest fringe of vegetation; 

Morinda citrifolia as bushes up to 10 ft high, mostly higher up the hillock from 
the fern tangle; 

Parinari corymbosa as a cluster of several much branched trees up to 20 ft high, 
at the top beside the Survey Beacon. 


I remember, but have failed to note, rather numerous large and metallic green 
‘rose-beetles’ buzzing about on the summit as if they were bees. On the west side of 
the summit, which was actually a slight ridge running north and south, there was a 
large nest of the sea-eagle, surrounded by fish-bones and the skeletons of sea- 
snakes. 


Half a mile north of P. Berhala, roughly in line with its slight ridge, there was a 
skerry of granite rocks just breaking the surface of the sea, about 40 yards across, 
and devoid both of sandstone and vegetation. It indicated another such islet worn 
down to the granite platform that was itself disintegrating. 


bE. ye stelle 


ee ae ee 
—— 


Fig. 6. Sketch of a transect across P. Berhala, with granite platform and sandstone hill. M, Morinda 
citrifolia; Pa, Parinari corymbosa; S, Stenochlaena. 


14 Gard. Bull. Sing. 38(1) (1985) 


Pulau Babi Tengah 


I visited this island off the Johore coast on 10 June 1932, 27 September 1936 and 
29 October 1936. It is a granite island with an accumulation of granite boulders on 
the points with sandy bays between them. It had been largely deforested but 
retained on the north and east shores a Terminalia-zone with its backing of Eugenia 
grandis, typical of the east coast of Johore. Coconut palms had been planted over 
most of the island. At the north-east end the small headlands or spits of granite 
boulders were connected with the main island by necks of small water-worn 
boulders and coral detritus on which Pemphis acidula formed elfin woods, almost 
pure stands, as at Tg Ruit. Between these spits, the sandy bays had small stands of 
mangrove as at P. Tulai. A sandy spit at the north-west end had a natural growth of 
Casuarina equisetifolia and some tufts of Spinifex littoreus on the growing front. 


It will be seen from the list in Table 1 that none of the plants characteristic of the 
granite islets of Tiuman occurred, unless the rather ubiquitous Atalantia and 
Diospyros. \ failed to record the ferns and coastal aroids and orchids. Hence there 
are many apparent absentees in Table 1, which account for its low total of species. 
It could have been that, before deforestation, the island had much of the flora of 
the granite islets off Tiuman. 


I make the following observations on a few of the plants:- Atalantia was very 
abundant. Barringtonia macrostachya and Garcinia nigrolineata were common 
inland. Pandanus odoratissimus var. laevis (P. spurius, as the thornless variety) 
grew under the coconut palms and might have been planted. The record of 
Schizachyrium sanguineum is the only one for the Malay Peninsula according to 
Gilliland. The shrub Timonius compressicaulis was common on rocky headlands on 
the north-east side of the island. Concerning Messerschmidia (Tournefortia) one 
would like to know its distribution in the Peninsula. 


Notes on special plants 


Allophylus cobbe — I made several varieties for this species in the Malay 
Peninsula (Corner 1939a). They have been recognised by Leenhouts (1967) who 
equates them as follows:— 


var. glaber as A. javanicus and A. longipes, 
var. limosus as near to A. ternatus, 

var. marinus as A. timorensis, 

var. velutinus as A. racemosus, 

var. villosus as A. villosus 


but he agrees with me that there is, practically, only one species. It would be 
interesting to pursue this problem experimentally, by raising seed of the varieties 
and by hybridisation. 


Breynia coronata ?, SFN 29853. — This was a straggling climber on P. Duchong, 
22 August 1935, common on rocks near the top of the island. The leaves were 
glaucous beneath. All the plants were sterile. 


Cissampelos ?, Corner s.n., P. Tulai, 18 August 1935; common climber in the 
Terminalia-zone, with yellowish leaves. 


Botany of eastern Pahang and Johore islets 15 


Calamus chibehensis, SFN 29842. — Furtado (1956) attributed this collection to 
M.R. Henderson, which is an error. I saw what I took to be the same plant also at 
P. Tulai and Juara Bay. It is probably no other than a variety of C. burkillianus 
which occurs on Tiuman. 


Celastrus ? SFN 29829, P. Chibeh, 19 August 1935. — This was a frequent small 
tree on rocks at the top of the island and on the west slope. It resembled the 
guava-tree (Psidium) in shape, leaf, and pale brown, slightly papery-flaky, bark. I 
collected it as Aporosa ?, but it was referred to Celastrus by M.R. Henderson. It is 
not listed by Ding Hou (1965). 


Chrysopogon fulvus. — This tussock grass occurs in Africa, India, and Thailand, 
but in Malaya it is known only from Kedah (Bukit Wang) and P. Tokong Burong 
(Gilliland 1971). It would seem to be carried somehow by terns, but that would not 
explain its strange distribution. 


Diospyros ferrea. — Concerning the variation in this very widespread species 
there is the article by Fosberg (1939). Recently, however, doubt has been thrown 
on the use of this name (Smith 1981). 


Ehretia sp., SFN 29827, P. Chibeh, 19 August 1935; SFN 29845, P. Sepoi, 20 
August 1935; straggling climber on more or less exposed rocks; berries small, dull 
orange. 


Ficus. — When I visited the islets, the identification of the strangling figs was in a 
state of confusion. I believe, now, that I mistook several records which I made on 
the spot as F. microcarpa. Certainly some, for which I made specimens, have 
turned out to be F. kurzii and F. stricta. Careful study of the venation of the dried 
leaves is necessary to identify these sterile saxicolous fig-plants. I note that F. 
hispida, abundant in the northern half of the Peninsula, was absent from the 
southerly islands, just as from the south of the Peninsula. 


Gnetum gnemon. — There are notes on the wild occurrence of this tree in Malaya 
(Corner 1939b). 


Hemiscolopia trimera. — The strange distribution of this small tree, resembling 
the rukam (Flacourtia), is given by Sleumer (1954), to whose account must be 
added his later records from Malacca and Ulu Sedili. It suggests a north-south line 
from Indo-China to the Sunda Straits. On P. Tulai it must be a relic. It may have 
had the wider distribution of Hydnocarpus ilicifolia in the northern part of the 
Peninsula from Langkawi across Kedah to Pahang. The tragedy is that with such 
intensive deforestation the Peninsula has lost much of its botanical history. 


Hydnocarpus ilicifolia, SFN 25762, P. Sepoi, 13 June 1932; SFN 29826, P. 
Chibeh, 19 August 1935. — Tree - 30 ft high; trunk cylindric or slightly fluted 
downwards; bark light brown to greyish brown, slightly flaky with thin angular 
pieces but appearing smooth, becoming shortly and finely fissured, not pustulate or 
rugose; inner bark pallid white, greenish below the outer bark; fruits 3-5 cm wide, 
subglobose, velvety black; seeds with thin oily sweet pulp round them. 


Manilkara kauki. — This tree, widely spread from Burma and Indo-China to 
north Australia, occurs in the Malay Peninsula only ‘on rocky headlands and 


16 Gard. Bull. Sing. 38(1) (1985) 


islands off the east coast of Johore and Pahang’ (Ng 1972). I found it only on P. 
Chibeh. Like Hydnocarpus ilicifolia, it seems to be a relic of the monsoon climate 
at the north of the Riouw Pocket (Corner 1978). 


Myristica guattertifolia. — The remarkable and almost coastal distribution of this 
‘very distinct species’ is given by Sinclair (1958, 1968). It is surely connected with 
the northerly shore-line of the Riouw Pocket as it impinged on the China Sea and 
extended along the east coast of Malaya to Sumatra and Java; in fact, it circums- 
cribes the Riouw Pocket. In Malaya it is known from P. Tenggol (Dungun), P. 
Tiuman, P. Chibeh, P. Setindan, and the coast of Johore from Mersing to the Sedili 
rivers (Corner 1978). 


Pandanus dubius. — As | have noted (Corner 1978), this tree-like species with 
stout stilt-roots and broad leaves reaches its western limit of distribution on P. 
Tenggol (Dungun), P. Tiuman and some of its islets, and Tanjong Sedili Kechil 
(Kecil). It is another indication of the northern limit of the Riouw Pocket. It is 
common on the rocks at Bako National Park in Sarawak where it is accompanied by 
the saxicolous state of Pandanus epiphyticus. That species occurs in the south-east 
of Malaya and indicates the more central part of the Riouw Pocket. Strangely, it 
seems never to occur on coastal rocks in the Peninsula (Corner 1978). 


Phyllanthus ?, SFN 29848, P. Sepoi, 20 August 1935; sprawling climber on rocks 
half-way up the island and at the south end; leaves subglaucous beneath; berries 
pinkish purple, small. 


Pisonia grandis. — This seashore tree, which I used to call P. excelsa, is widely 
distributed on rocky coasts in the west Pacific but, in Malaya, only on the granite 
islets about Tiuman and at Kuala Trengganu. Its sticky fruits adhere to the feathers 
of birds which distribute it, and it is thought to succeed only in soil enriched with 
guano. It appears, like Pandanus dubius, to have come westwards; yet, unlike that 
pandan, it has not been recorded from the China Sea coast of Borneo. There is a 
photo of it, which I took on P. Sepoi, in the Flora Malesiana (Stemmerik 1964). 
Concerning the general occurrence of Pisonia forest and its competition with 
Ochrosia oppositifolia, there is the article by Fosberg (1976). 


Planchonella firma. — This is said to be a tree of mossy mountain forest in 
Malaya, with P. Setindan as the only lowland record (Ng 1972). There are other 
plants, such as Baeckia, Styphelium, and Vaccinium bracteatum, which move from 
mountain to lowland, but none of these occurred in the islets around Tiuman. It 
may be a relic of the mountain flora of P. Aor and P. Tinggi, and it should be 
looked for on Gunong Panti. See, also, the following note. 


Planchonella linggensis. — This species is distributed throughout Malesia to 
Australia and the Pacific islands, but in the Malay Peninsula it appears to be 
restricted to the rocky sea-coast of Penang, Pahang, and east Johore, except for 
records from Penang Hill and Mt Ophir (Ng 1972). I found it on P. Chibeh, P. 
Tulai, and P. Duchong. It is a remarkable relict occurrence. 


Polyalthia sclerophylla. — Concerning the identity of the collection from P. 
Duchong, I have already published a note (Corner 1978). 


Premna obtusifolia. — This name supplants those of P. corymbosa and P. 
integrifolia, which have been widely used for this common plant (Fosberg 1953). 


Botany of eastern Pahang and Johore islets 17 


Tristania obovata — Tanjong Penyabong and P. Setindan appear to be the 
northern limits, on the east coast of Malaya, of this species of the Riouw 
Archipelago. 


Table 1. List of flowering plants and ferns at Tanjong Ruit (R), Pulau Chibeh (C). 
P. Sepoi (S), P. Duchong (D), P. Setindan (St), and P. Babi Tengah (B). 


(+ means present; — means absent or not found) 


BR mGalc§ns Dri Sou.B 


TREES. SHRUBS. PALMS. PANDANS 


Adenanthera pavonina ee ee ee, ee 
Allophylus cobbe v. limosus 25. SPs yolhss wicetoin¥ 

V. marinus +. (ee SASS Hee Oo 

v. velutinus en ene Se 
Antidesma cuspidatum nai MATER ier Sols 
Ardisia crispa 6 See ee ee 
Ardisia elliptica oo a ee ee on ee 
Callicarpa longifolia eth A? egtion gage 
Caryota mitis Ee San Se ee 
Casuarina equisetifolia Ses ofclianit apes el 
Cedrela ? fe ae ee 
Cerbera manghas - =  =eeeietes iis 
Chionanthus ramiflorus — so epighaatn ides 3 


Cocculus ovalifolius eS 


18 


Trees, etc., cont. 


Cordia subcordata 
Croton heterocarpus 
Cycas rumphi 
Cynometra ramiflora 
Decaspermum paniculatum 
Desmodium umbellatum 
Diospyros ferrea 
Diplospora malaccensis 
Dracaena maingayl 
Elaeocarpus floribundus 
Erioglossum rubiginosum 
Erythrina indica 
Erythroxylon cuneatum 
Eugenia claviflora 
Eugenia glauca 

Eugenia grandis 
Eugenia longiflora 
Eugenia palembanica 
Eugenia polita 

Eugenia subdecussata 
Eurycoma longifolia 
Excoecaria agallocha 
Ficus caulocarpa 

Ficus crassiramea 

Ficus deltoidea 


Ficus drupacea 


Gard. Bull. Sing. 38(1) (1985) 


- =~ = + 
._ a 
- = = + 
in fact ee 
- + + +4 
- = = + 
yer re 
=, th~ .e 
Seyuey 1978}.- 
=. aaah olla eee 
- - + = + 
+ = + $+ = 
Lie aah Ulpetanaai 
2 vena Ay ee 
+ = + + + 
en 
meeer ee te 
x. eheiuely ene a 
1. «nda 
wee 
L appa 


Botany of eastern Pahang and Johore islets 


Trees, etc., cont. 


Ficus fistulosa 

Ficus grossularioides 
Ficus hispida 

Ficus kurzii 

Ficus microcarpa 

Ficus stricta 

F. sundaica v. beccariana 
Ficus superba 

F. tinctoria ssp. gibbosa 
Ficus variegata 

Garcinia hombroniana 
Garcinia nigrolineata 
Gelonium glomerulatum 
Glochidion littorale 
Gnetum gnemon 

Grewia paniculata 
Guettarda speciosa 
Guioa pleuropteris 
Heritiera littoralis 
Hernandia nymphaeifolia 
Hibiscus tiliaceus 
Hydnocarpus ilicifolia 
Ixora congesta 
Kleinhovia hospita 


Knema globularia 


St 


19 


20 


Trees, etc., cont. 


Knema malayana 

Leea indica 

Lepisanthes fruticosa 
Litsea glutinosa 
Macaranga javanica 
Mallotus tiliaefolia 
Manilkara kauki 
Melastoma sanguineum 
Memecylon coeruleum 
Memecylon myrsinoides 
Memecylon ovatum 
Messerschmidia argentea 
Morinda citrifolia 
Myristica guatteriifolia 
Myrsine porteriana 
Neolitsea zeylanica 


Ochrosia oppositifolia 


Gard. Bull. Sing. 38(1) (1985) | 


Oncosperma filamentosa 


Pandanus dubius 


Pandanus odoratissimus 


Parinari corymbosa 
Pemphis acidula 
Phoebe declinata 


Pisonia grandis 


Pithecellobium contortum 


Pithecellobium ellipticum 


= Jyh 
ashivitn hee 
= 2 yyeeee 
— 
= + ,obneae 
=. Syn 
+ + + = 
=  —?wieseaueane 
nonoleseee 
- = = + 
- + + + 
a\3 weil wel 
= ete eee 
= yin 
yo ee 
- + + + 
4: yabaiihaeotel 
- = = + 
=  yriopmepete 
so\iqaed eet 
~ vinokerdolaantins 
ao. 


Botany of eastern Pahang and Johore islets 


Trees, etc., cont. 


Pittosporum ferrugineum 
Planchonella firma 
Planchonella linggensis 
Planchonella obovata 
Podocarpus polystachyus 
Polyalthia sclerophylla 
Pongamia pinnata 
Premna obtusifolia 
Scaevola taccada 
Sterculia foetida 
Stereospermum fimbriatum 
Terminalia catappa 
Thespesia populnea 
Timonius compressicaulis 
Trema amboinensis 
Trigonopleura ? 
Tristania obovata 

Vitex pubescens 
Wendlandia ternifolia 
CLIMBERS 

Aganosma marginata 
Caesalpinia bondhuc 
Caesalpinia crista 
Calamus chibehensis 
Calamus perakensis 


Canthium confertum 


St 


21 


2? 


Climbers cont. 


Cassytha filiformis 

Cissus repenns 

Comsants monospermus Vv. malayanus 
Derris scandens 

Derris thyrsiflora 

Derris uliginosa 

Ehretia SFN 29827, 29845 
Ficus parietalis 
Flagellaria indica 
Gnetum latifolium v. funiculare 
Gynochthodes coriacea 
Hoya diversifolia 
Ipomoea illustris 
Loeseneriella pauciflora 
Morinda umbellata 
Paramignya andamanica 
Parsonsia spiralis 
Phyllanthus ?:‘SFN 29848 
Pisonia aculeata 
Psychotria sarmentosa 
Salacia chinensis 
Sauropus albicans 
Sauropus sp. (s.n.) 
Schefflera venulosa 
Tetracera assa 


Tristellateia australasica 


Gard. Bull. Sing. 38(1) (1985) 


- - = + 
eo 
xy vodey oleae 
von ig Se 
+ + + = 
+ xeon eee 
 i\ cipal 
2” oka ei 
_- 2s eee 
Aout) ate 
- + + + 
a. yell aa 
oi 7 pral cee 
See 
tj ee 
4. cynical 
a a 
aside lean 
+ £f + = 
<viiveieacee 
- + + + 


Botany of eastern Pahang and Johore islets 


Climbers cont. 

Vitis japonica 

Yellow, cordate-leafed climber 
HERBS, SEDGES, GRASSES 
Aneilema sp. SFN 29794 
Bulbophyllum vaginatum 
Canavalia turgida 
Cansjera zizyphoides 
Chasalia curviflora 
Coelorrachis glandulosa 
Commersonia platyphylla 
Cyperus cyperinus 
Cyperus diffusus 
Cyperus dubius 

Cyperus javanicus 
Cyperus kyllingii 
Cyperus radians 

Dianella ensifolia 
Digitaria ciliaris 

Eulalia ridleyi 
Fimbristylis dichotoma 
Hedyotis verticillata 
Ipomoea pes-caprae 
Ischaemum muticum 
Lasianthus cyanocarpus 
Oldenlandia paniculata 


Panicum repens 


St 


23 


24 


Herbs, etc., cont. 


Paspalum conjugatum 
Plocoglottis porphyrophylla 
Pseuderanthemum crenulatum 
Schizachyrium sanguineum 
Scleria lithosperma 

Spinifex littoreus 
Sporobolus virginicus 
Taeniophyllum serrula 
Themeda villosa 

’ Vandellia crustacea 
Vandellia hirsuta 

Vigna marina 

Wedelia biflora 

Zoysia matrella 

EPIPHYTES ON EXPOSED ROCKS 
Asplenium nidus-avis 
Cymbidium finlaysonianum 
Davallia solida 
Dendrobium crumenatum 
Dendrobium serra 
Dischidia rafflesiana 
Drynaria quercifolia 
Fagraea auriculata 
Hydnophytum formicarium 
Medinilla hasseltii 


Polypodium scolopendria 


Gard. Bull. Sing. 38(1) (1985) 


= -b <= = 
—— = 4 2 
- = = + 
- = = + 
—_ oe = a 
ws — =< = 
= oe = = 
- + + + 
che 2s + = 
- + + + 
- + + + 
as — + ag 
- = + + 
— = + — 
aed <= 4 = 
as + = = 
+ + + = 


Botany of eastern Pahang and Johore islets 25 


Epiphytes cont. RovoC S DH Seni B 
Scindapsus sp. eee, Seer se a 
SFN 29816 : eee era ae 


FERNS (OTHER THAN EPIPHYTES) 


Adiantum stenochlamys Bue) Re aC tigen 
Asplenium glaucophyllum 2B eee eae = a 
Asplenium macrophyllum ED 30, SER ee 2 Se a 
Blechnum orientale ES ees oe ee 
Cyclophorus adnascens Ss ee i 
Lygodium flexuosum Ppa 2 C2. a/R i a 

~ Nephrolepis biserrata 2 Ur: are Rens Set dee 
Pteridium aquilinum aS gine by aes Came © VC as ee 
Pteris ensiformis EE pee + nee oe 
Stenochlaena palustris ts Ay ee 5 me tag et ale 
Vittaria elongata SSA Sears a en 
Reem et ee Ge. 65 83-94 40 

References 


Corner, E.J.H. (1939a). Notes on the systematy and distribution of Malayan 
phanerogams, I. Gdns’ Bull, Singapore 10: 1-55. 


. (1939b). Notes on the systematy and distribution of Malayan phanero- 
gams, III. Gdns’ Bull. Singapore 10: 239-329. 


. (1978). The Freshwater Swamp-forest of South Johore and Singapore. 
Gdns’ Bull. Singapore, Suppl. n. 1. 


Ding Hou (1965). Celastraceae. Identification List 24. Flora Malesiana Foundation, 
Leiden. 


Fosberg, F.R. (1939). Diospyros ferrea (Ebenaceae) in Hawaii. Occas. Papers 
Bernice P. Bishop Mus., Hawaii, 15 n. 10: 119-131. 


. (1953). Nomenclature of Premna obtusifolia R. Br. Taxon 2: 88-89. 


. (1976). Coral island vegetation. Biology and Geology of Coral Reefs vol. 
III, Biology 2: 256-277. Academic Press Inc.: New York, London. 


%6 Gard. Bull. Sing. 38(1) (1985) 


Gilliland, H.B. (1971). Grasses of Malaya. Revised Flora of Malaya vol. III. 
Singapore Botanic Gardens. 


Leenhouts, P.W. (1967). A conspectus of the genus Allophylus (Sapindaceae). 
Blumea 15: 301-358. 


Ng, F.S.P. (1972). Sapotaceae. Tree Flora of Malaya vol. I. Longman Group Ltd. 
London. 


Sinclair, J. (1958). A revision of the Malayan Myristicaceae. Gdns’ Bull. Singapore 
16: 205-472. 


. (1968). Florae Malesianae Praecursores — XLII. The genus Myristica in 
Malesia and outside Malesia. Gdns’ Bull. Singapore 23: 1-540. 


Smith, A.C. (1981)). Flora Vitiensis Nova vol. 2. Pacific Tropical Botanical Gar- 
den: Kawai, Hawaii. 


Stemmerik, J.F. (1964). Nyctaginaceae. Flora Malesiana ser. 1, 6: 464. 


‘Ol 94] UO DIUadNT pure VIPVUIUAA] JO \SIIOJ [PISPOD 


oy) pur ‘osjuso oy) Ul srydudag YW SNUY)SI AYSOI OY) “JYSII OY) UO SI puR[pRoY YIPUIWAA | OUT 
‘TEH] Ul SuPUNTy Neng ‘ying dsuoluey “| eI “ 


‘CEOL 


‘uRWINIT “dg ‘NY 3] JO snumyjst AYO dy} UO BjNpiIv siydwag 


C Meld 


28 


Plate 3. Pulau Chibeh from the south-west, 1935. 


Plate 4. P. Chibeh, north end, 1935. 


ME hee 


Plate 5. P. Chibeh from the north-east, 1935. 


Plate 6. P. Chibeh, rocks at the south end, with the plant-collector Kiah bin Mohd. Salleh; 1935. 


30 


‘TEST “You ay Wor 1odag neing “/ a1eIg 


31 


Pi Measksncmipamastcmce 


Weare 


oe eae e 


ope 


Plate 8. P. Sepoi from the east, 1932. 


% 


Plate 9. P. Sepoi at the north-west end, 1935. 


32 


“SOYOURIG “IIL JSOWR “ITYM YIM SIPUDAS DIUOSI JO SId1 J. 
‘SE6l ‘Pud Jsvd-YINOS dYI Iv IOdI|Sg “g 


‘OT eld 


co 
or 


“SyYUNI}-991] pue SIOp[nog dy} SuIdseyd s}OOI-sIJ Jo ajsue} dU]. 
“SGI “ISA10} ay} UI 1odag “gq 


‘TT Id 


34 


35: 


de the forest, 19 


1 1nsl 


P. Sepo 


Plate 12. 


oe) 
Oy 


The fluted trunks of Atalantia monophylla. among the fig-roots (mainly of Ficus superba). 


The plant-collector Kiah bin Mohd Salleh in the middle distance to the right. 


36 


a, 


® 


naige 


1933: 


Mohd Salleh 


in 


ith undergrowth of Stenochlaena palustris and the 


is W 
ant-collector Kiah b 


isonia grandi. 
the pl 


. 
| 


P. Sepoi in the forest of P 
Atalantia-forest behind 


Plate 13. 


[SI URL SY} JO JUWUWINS 9Y} WOIJ UDIS se spUP]sI []eUIS OM) OUT, 
uoINg SUOYO] Neng ‘pl Meld 


“SC6l ‘SuoINng 
SUOYOL “d JO puvjsi urew sy) Jo doy ayi uo snaynf uoSodosduyD sse13 dy) JO sydossnq ay]. “ST eld 


‘VJO] 94) 0} ‘pue 


‘SC6l “DIUO]sUlDg 
DIYUIUAIT JO IBLIUOIJ JY} BUIMOYS ‘jsed 94} WOIJ ne] BuOyoONG Neng 


‘OL Id 


39 


oe, 


‘ZEST SONSIMUIG “AA “YB ay) UO andy 
pods SY} {90UR}SIP 94} UI eTRD LS youne] sy} YIM ‘WOH ed o}1UeIs oY} SuIMOYs ‘eTeYyIIg “dq 


“LI Id 


40 


‘TET -3eY 94) UO 
IPPYOOI P SE BUIMOYS DSOqUIA10) LIDULAD JO SUMOID JY} YIM ‘YOU dy) Wo E[RYIOG Neng ‘RT WeId 


Plate 19. 
‘The rocks to the north 
of P. Berhala; 1932. 


Plate 20. 

P. Berhala, the 
granite platform 
with the base of the 


right, and in the 
distance the low 
rocky island to the 
north; 1932. 


Plate:21; | 
P. Berhala, showing: 
the granite platform 
from the summit of 


the hill; 1932. 


New Species of the Genus Homalomena (Araceae) from Sumatra with 
a Short Note on the Genus Furtadoa 


Mitsuru HOTTA 


Biological Laboratory, Yoshida College 
Kyoto University, Kyoto 606, Japan 


EFFECTIVE PUBLICATION DATE: 31 AUG. 1985 


Abstract 


Five new species of the genus Homalomena (Araceae) from Sumatra are described and their 
relationships and chromosome numbers briefly discussed. Among them, Homalomena rusdii sp. nov. 
stands out by its free ligule at the petiole base and by its (usual) basal placentation of ovules. 
Homalomena mixta is transferred to the genus Furtadoa on the basis of the floral characters. 


From 1980 to 1984, I had several opportunities to visit West Sumatra as a 
member of the Sumatra Nature Study (SNS) Project, cooperating in field studies of 
Andalas University (Indonesia) and Kyoto University (Japan). Members of the 
Botany Group of the Project had been gathering many herbarium specimens 
including a rich and good aroid collection (Hotta, 1984), and during those travels I 
had opportunities to work on the taxonomy of Sumatran flora at the Herbarium 
Bogoriense and the Singapore Botanic Gardens’ Herbarium. 


The aroid flora of Sumatra is characterized by a slightly poorer diversity than that 
of Borneo, especially among those elements that have a limited distribution in the 
wet tropical region of Malesia. Borneo has 6 endemic genera of Araceae: Arid- 
arum, Bucephalandra, Heteroaridarum, Hottarum, Pedicellar'um and Phymatarum 
but Sumatra does not have endemics since Furtadoa once believed to be endemic to 
Sumatra, has now been found in Malaya (see end of article). On the other hand, 
in Sumatra, the genus Homalomena is rich in species and shows much variation. In 
this paper, new and interesting species from our Sumatran material of Homa- 
lomena are described. 


Homalomena megalophylla M. Hotta, sp. nov. (sect. Homalomena) Fig. 1,2 &4A 


Herba maxima caudiculo erecto 0.5-1.5 m longo, 8-15 cm crasso. Foliorum petiolus crassiuscule quam 
lamina 2-plo longior, 1-1.6 m longus, ad 13-4, longitudinis vaginatus, lamina subcoriacea, supra obscure 
viridis, subtus pallide viridis, ovato cordata, 50-80 cm longa, 35-50 cm lata, lobis posticis rotundato- 
oblongis usque 20 cm longis, lobo antico ovato, acuminato, nervis lateralibus I. utrinque cire. 3 
basalibus, 5-6 costalibus adscendentibus prope marginem sursum curvis. Pedunculi plures usque 25-30 
cm longi. Spathae pars inferior oblongo-ovoidea (4)5-6.5 cm longa, 1.5-2.5 cm ampla, pars superior 
convoluta 7-10 cm longa. Spadicis stipite 3-5 mm suffulti inflorescentia feminea (2.5)3.5-4(4.8) cm 
longa, 1.5(-2) em crassa, mascula 6.5-8 cm longa. Flores masculi 4-5(-6) andri. Pistilla oblongo- 
obovoidea virescentia, stigmate discoideo instracta; ovaria inferne trilocularia, placentis 3 in quoque 
loculo a centro prominentibus, superne unilocularia, placentis parietalibus, ovulis numerosis alfixa. 
Staminodia claviformia. Baccae obovoidea 7 mm longa; semina ellipsoidea 1.5 mm longa. 


WEST SUMATRA: Airsirah pass, 850-900 m, July 28, 1984, M. Hotta, H. Okada & T. Kohyama 8 
(KYO); between Sungai Dareh and Sijunjung, steep-rocky cliff off the road side, alt. 100-200 m, Aug. 
20, 1981, M. Hotta, H. Okada & R. Tamin 104 (Holotypus in KYO, isotypus in BO); Harau, 
Pajakumbuh, rocky open place at the foot of the hill, alt. 600 m, Aug. 27, 1983, M. Hotta & R. Tamin 
299 (KYO). 


43 


M.Umebayash 


Fig. 1. Homalomena megalophylla M. Hotta 
Leaf, X *%; and habit, x Mo. 


44 


Fig. 2. Homalomena megalophylla M. Hotta 
A: spathe (right) and fruiting spadix (left), x 12; B: fruit (right) and its longitudinal section 
(left), X 5; C: cross-sections of upper (above) and lower (below) parts of ovary, x 5; D: 
close-ups of two male flowers, front view (right) and a stamen, side view (left), x 10; E: seed, x 
10. 


This new species is one of the largest plants found among the species of the genus 
Homalomena. It seems to be related to H. rubescens (Roxb.) Kunth, the rela- 
tionship being indicated by the male flower, which has large ovoid pollen sacs 
attached directly to the surface of the spadix axis (cf. Engler 1912: p. 64, fig. 39, D 
& E). However, my new species is distinctively different in that it has a constricted 
spathe and is an extremely large plant. In the latter respect it is like H. gigantea 
Engler but differs from that by the shape of the stamen and that of the spathe. 


H. megalophylla is a common and conspicuous aroid on open, wet, rocky slopes 
in West Sumatra. It is therefore strange that specimens are not found in the Bogor 
or Singapore herbaria. The chromosome number of this species is 2n=40 (based on 
counts in Airsirah clones made by H. Okada, unpublished). 


Homalomena gadutensis M. Hotta, sp. nov. (sect. Homalomena). Fig. 3,4D 


Herba majuscula caudiculo erecto 10-40 cm longo, 1.5-2 cm crasso. Foliorum petiolus quam lamina 
1.5-2-plo longior, 30-50 cm longus, ad 13-2 longitudinis vaginatus, lamina supra viridis, subtus pallide 
viridis, ovato cordata, 13-25 cm longa, 10-15 cm lata, lobis posticis ovato-triangularis usque 5 cm longis, 
lobo antico ovato, acuminato, nervis lateralibus I. utrinque circ. 3 basalibus, 5-6 costalibus adscendenti- 


45 


Fig. 3. 


46 


Homalomena gadutensis M. Hotta 

A. habit, x 2; B: spathe (left) and spadix (right), x */4; C: close-up of female part of spadix, x 
7; D: close-up of male part of spadix, x 7; E: female flower (/eft) and staminode (right), « 9; F: 
longitudinal section of ovary, X 9; G: cross-section of ovary, X 15; H: ovule, x 40; J: side view 
(above) and longitudinal section of male flower (below), x 10. 


Homalomena and Furtadoa 47 


bus prope marginem sursum curvis. Peduncli plures usque 8-15 cm longi. Spathae pars inferior oblongo- 
ovoidea 2.5-3.8 cm longa, 1.2-1.5 cm ampla, pars superior naviculiformia, 5-7.5 cm longa. Spadicis 
stipite 10 mm suffulti inflorescentia feminea 2-2.5 cm longa, 0.5-0.6 cm crassa, mascula 4.5-6 cm longa. 
Flores masculi (3-)4(-5) andri. Pistilla oblongo-obovoidea virescentia, stigmate discoideo instracta; 
ovaria inferne trilocularia, placentis 3 in quoque parietalibus, ovulis numerosis affixa. Staminodia 
superne obconica, inferne filiformia. 


WEST SUMATRA: Ulu Gadut, in forest floor near hill ridge, alt. 500 m, Nov. 29, 1980, M. Hotta 
25091 (KYO, BO), alt. 550 m, Nov. 29, 1980, H. Hotta 25105 (Holotypus in KYO, isotypus in BO); 
Pinang Pinang, Ulu Gadut, common on forest floor, alt. 400-800 m, Dec. 17, 1982, M. Hotta, H. Okada 
& M. Ito 88 (KYO); G. Kambot, alt. 400-650 m, Jan. 23, 1981, M. Hotta & R. Tamin 255 (KYO), 262-b 
(KYO), 270 (KYO); Gajabuih, common on forest floor, alt. 400-650 m, Jan. 5, 1981, M. Hotta 25830 
(KYO, chromosome number 2n=38, cited by Dr. H. Okada, 1984 as Homalomena sp. nov. 1), 450-700 
m, July 31, 1984, M. Hotta, H. Okada & T. Kohyama 101 (KYO); Batu Bajolang, forest floor near 
ridge, alt. 400-600 m, M. Hotta, H. Okada & M. Ito 1132 (KYO, BO, AND%). 


Fig. 4. Leaf shape variation of allied species of Homalomena in Mt. Gadut area 
A: H. megalophylla; B: H. sagittifolia; C: H. pendula; D: H. gadutensis. 


H. gadutensis is closely related to H. sagittifolia, one of the very variable species 
in Malesia. Collections of the latter related species in the same group (H. pendula 
group, H. megalophylla and H. gadutensis) from Mt. Gadut area vary a great deal 
in shape and size of the leaves (Fig. 4), and in the spathe (Fig. 5). Precise 
identification of each species in the group could not have been achieved without 
examining the characters of the spathe and the male flower. The leaf size shows a 
tendency to decrease as the elevation increases. This tendency is illustrated by 
measurements of the largest leaf of each collection, after classification using spathe 


*AND = Herbarium of Andalas University, Ulu Gadut, Padang, West Sumatra, Indonesia. 


G. Kambot 


Nn 
6 5 
B 4 
a 
Oe 
o4 
a. 
S 
oO 
2 
foe) 
g Ulu Gadut 
ie) 
& 5 
n 4 
= 3 
2 
as) 1 
oO 
oO. 
Nn 
Lol 
) 
7 Karang Puteh 
© 6 
= 5 
34 
Z 3 
2 
1 
0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 


Ratio of spathe-lamina length to total spathe length 


Fig. 5. Variation of spathes of four Homalomena taxa in Mt. Gadut area 
Ratio of the length of spathe lamina to the total length of spathe (horizontal axis), and number 
of specimens from three local populations (vertical axis); ratio 0 = H. pendula group (spathe 
without constriction and undifferentiated into the basal (=tube) part and apical lamina), 
02-0.6 = H. sagittifolia, and over 0.6 = H. gadutensis. 


characters (see Fig. 5), i.e., (a) spathe unconstricted — H. pendula species group, 
(b) spathe constricted and separable into the spathe tube and the lamina. Group (b) 
is subdivided into (1) ratio of lamina/total length of spathe 0.2-0.6 — H. sagittifolia, 
and (2) ratio of lamina/total length of spathe 0.6 or more — H. gadutensis. From 
the analysis, it is seen that H. gadutensis also has smaller leaves and is a compara- 
tively smaller plant. Besides, the lamina of its spathe is white and boat-shaped, 
spreading open widely at the time of flowering. The chromosome counts for H. 
pendula group, H. megalophylla and H. sagittifolia of Mt. Gadut area are in all 
cases 2n=40 or 80, but is 2n=38 for H. gadutensis, a number peculiar for the genus 
(H. Okada 1984 & unpublished). 


A variety, H. sagittifolia var. sumatrana v. Ard. v. Rosen., may at first seem to 
be H. gadutensis on account of the smaller size of the entire plant and its occurrence 
in a locality at a higher altitude (Talu, Ophir, 950 m, Biinnemeijer 1299, lectotype 
in BO), but this specimen, to my mind, represents a mountain ecotype of H. 
sagittifolia with a spathe-lamina which is rather short; and furthermore, H. sagittifo- 
lia occasionally does occur on mountains above 1000 m in West Sumatra. This 
mountain ecotype usually has a tetraploid chromosome complement, 2n=80, cited 
by Dr. H. Okada (unpublished). 


Homalomena padangensis M. Hotta, sp. nov. (sect. Chamaecladon) Fig. 6 


Herba majuscula rhizomate obliquo 3-8 cm longo, 1-1.5 cm crasso, pauce (2-4) foliato. Foliorum 
petiolus quam lamina 1.5-2-plo longior, 30-40 cm longus, ad % longitudinis vaginatus, lamina sub- 
coriacea, supra vertinus nitida, viridis, subtus pallide viridis, cordata, paullum inaequilatera, 15-25 cm 
longa, 10-15 cm lata, apice breviter acuminata, nervis lateralibus I. utrinque 5-8 adscendentibus leviter 
arcuatis, nervis secundariis inter primarios numerosis interjectis. Pedunculi usque 3-4 cm longi. Spathae 
oblonga breviter apiculata, viridis, 2 cm longa, 6-8 mm ampla. Spadicis sessilis oblongi inflorescentia 
feminea 4 mm longa, quam mascula 4-plo brevior. Flores masculi (1-)3 andri. Pistilla ovoidea virescen- 


tia, stigmate discoideo coronata; ovaria bilocularia, ovulis medio affixis. Staminodia crassa claviformia 
quam ovaria duplo brevior. 


48 


Fig. 6 Homalomena padangensis M. Hotta 


A: habit, x 42; B: spathes x 1; C: spadix, x 1; D: male flower, front (below) and side views 


(above) X 12; E: cross-section (lower left) and longitudinal (lower right) sections of female 
flower with staminode (above), x 8. 


49 


50 Gard. Bull. Sing. 38(1) (1985) 


WEST SUMATRA: Karang Puteh, on wet and open limestone cliff, alt. 150-300 m, Feb. 16, 1981, M. 
Hotta & H. Okada 344 (KYO, chromosome number 2n=40, cited by H. Okada 1984 as Homalomena 
sp. nov. 2), alt. 100-200 m, Aug. 14, 1981, M. Hotta & H. Okada 476 (Holotypus in KYO, isotypus in 
BOD, AND, SING & L). 


H. padangensis seems to be related to H. griffithii, a common and widely 
distributed species in western Malesia, but distinctly differs from the latter by its 
coriaceous leaf with a velvet-like gloss, cordate leaf base, and the decumbent stem 
with many strong roots growing out from the nodes.. H. padangensis is characte- 
rized by the very short filament of the stamen of the male flower (the pollen sac 
being attached directly to the spadix axis), and the relatively fewer female flowers 
in the lower part of the inflorescence. This species occurs in a limited area on the 
limestone hill of Karang Puteh and needs careful protection. 


Homalomena hastata M. Hotta, sp. nov. (sect. Chamaecladon) Fig. 7 


Herba majuscula caudiculo obliquo 3-5 cm longo, 1 cm crasso, 1-2 foliato. Foliorum petiolus laminae 
subaequilongus vel ea 2-plo longior, 40-60 cm longus, vagina 4-5 cm longa instructus, lamina supra 
opaca viridis, subtus pallide viridis, triangulari-oblonga inaequilatera, leviter curvata, basi hastata, apice 
acuminata, 25-35 cm longa, 10-15 cm lata, lobis posticis triangularis usque 5-8 cm longis, nervis 
lateralibus I. utrinque circ. 2 basalibus, 4-5 costalibus adscendentibus. Pedunculi usque 5-6 cm longi. 
Spathae ovoidea 2 cm longa, 0.8 cm ampla, apice cuspidata. Spadicis sessilis inflorescentia feminea 0.7 
cm longa, 0.4 cm crassa, mascula 0.8 cm longa. Flores masculi 3 andri. Pistilla oblongo-obovoidea 
virescentia, stilo cylindrici et stigmate discoideo instracta; ovaria inferne trilocularia, placentis 3 in 
quoque locullo a centro prominentibus, superne unilocularia, placentis parietalibus, ovulis numerosis 
affixa. Staminodia elongato claviformia inferne filiformia, pistilla aequantia. 


WEST SUMATRA: Along road between Lubuksikaping and Bondjor, Lurah Berangin Nature 
Reserve, 400-500 m, Sept. 21, 1984, M. Hotta 30231 (KYO); Pinang Pinang plot, Ulu Gadut, in dense 
forest floor of limestone area, alt. 500 m, Aug. 25, 1981, M. Hotta & H. Okada 624 (KYO), Aug. 27, 
1981, M. Hotta & H. Okada 689 (KYO), Sept. 2, 1981, M. Hotta 26670 (Holotypus in KYO, isotypus in 
BO),, Feb. 6, 1983, M. Hotta, H. Okada & M. Ito 1082 (KYO, BO); upper part of Pinang Pinang ridge, 
700 m, Aug. 1, 1984, M. Hotta, H. Okada & T. Kohyama 221 (KYO); G. Kambot, alt. 400-600 m, Jan. 
23, 1981, M. Hotta & R. Tamin 269 & 291 (KYO). 


In general H. hastata can be regarded as a plant well adapted to the habitat on 
the forest-floor. There are few leaves (usually 1 to 2) with thin and dark green 
lamina. The petiole is slender and very shortly sheathed at the base. The stem is 
somewhat decumbent and branches underground. The hastate leaf blade and the 
long filament of the male flower, both distinctive characters for the section Cha- 
maecladon, are also found in this species. The chromosome number of this species 
is 2n=40 based on a clone of Pinang Pinang ridge (H. Okada, unpublished). 


This new species occurs commonly in and around the Pinang Pinang Plot (one of 
our study sites for forest ecology on the gentle ridge of Pinang Pinang Hill), and we 
also found it in a few other places of the Mt. Gadut area and once in northern W. 
Sumatra (Lurah Berangin). It seems to have a limited distribution in and around 
the limestone areas of West Sumatra. 


Homalomena rusdii M. Hotta, sp. nov. (sect. Chamaecladon) Fig. 8 


Herba parvula, rhizomate repente, 3-10 cm longo, 1-1.5 cm crasso, dense foliato. Foliorum petiolus 
quam lamina 1.5-plo longior vel ei aequilongus, 8-13 cm longus, basi late vaginatus, vagina ovato- 
triangulare (2.5-3 cm) liguliformi-producta, lamina coriacea, supra nitida viridis, subtus pallidior, 
lanceolata vel oblongo-lanceolata, 8-11 cm longa, 2-3.5 cm lata, apiculo 2 mm longo cylindriformi, 
nervis lateralibus I. utrinque 3-4 angulo acuto adscendentibus leviter arcuatis. Pedunculi tenui usque 4-5 
cm longi. Spathae oblonga breviter apiculata, viridis, 1.5-2 cm longa, 6-8 mm ampla. Spadicis sessilis 


Fig. 7. 


Homalomena hastata M. Hotta 

A: habit, x 2 (Hotta, Okada & Ito 1082); B: spathe (left) and spadix (right), 1%; C: fruiting 
inflorescences, X 2 (Hotta & Tamin 291); D: close-up of female part of spadix, x 10; E: 
close-up of male part of spadix, x 10; F: female flower with staminode, x 15; G: staminode 
from base of spadix, x 15; H: longitudinal section of ovary (left), cross section of style (right), 
x 15; J: cross-sections of ovary at upper (left) and lower (right) parts, x 15; J: ovule (right) and 
seed, X 40 (left); K: male flower (side view), x 15; L: stamen, x 15. 


51 


Fig. 8. 


52 


Homalomena rusdii M. Hotta 

A: habit, x 2; B: leaf apices, x 4; C: petiole bases with free ligule, x 112; D: inflorescences, x 
2; E: spathe (left) and spadix (right), x 1; F: female part of spadix, x 8; G: longitudinal 
sections of 3 ovaries, X 13; H: ovule, x 40; J: stamens (front view), J: stamen (side view), X 40; 
and K: male flower (front view), x 40. 


Homalomena and Furtadoa 53 


oblongi inflorescentia feminea 2 mm longa, quam mascula 5-plo brevior. Flores masculi (1-)3 andri. 
Pistilla late ovoidea virescentia, in stilum brevissimum stigmate discoideo instructa; ovarium unilocu- 
lare, placentis 1, ovulis hemianatropis medio vel basi-affixis. Staminodia claviformia quam ovaria duplo 
breviora. 


WEST SUMATRA: Fort de Kock, Harau, Aug. 5, 1931, Frey Wyssling 154 (BO); Arau (Harau) 
Nature Reserve, Pajakumbuh, on wet rock near waterfall, 600 m, Aug. 27, 1983, M. Hotta & R. Tamin 
300 (Holotypus in KYO, isotypus in BO & L). 


H. rusdii has the distinct characters of the genus Homalomena, such as the free 
ligule of the petiole-sheath, and a uniloculate ovary with parietal-basal placenta- 
tion. It is undoubtedly also closely related to H. paucinervia by the coriaceous, 
lanceolate leaf blade with a few primary lateral veins (usually 3-4 on each side). H. 
paucinervia is a typical rheophytic aroid which is widely distributed in West Malesia 
(Borneo, Malay Peninsula and Sumatra), and two Sarawak collections (Hirano & 
Hotta 1284, and Hotta 15380) and one from southern Thailand (collection of Dr. T. 
Yahara and cultivated in the green-house of Kyoto University) have uniloculate 
ovaries with basal placentation. This type of unicarpellate ovary in Homalomena 
with parietal or basal placentation, has been reported on in H. minutissima (Hotta 
1967), closely related to H. humilis that has no direct relationship with the present 
species. The peculiar character of the uniloculate ovary in Homalomena might have 
been evolved in a parallel way within two groups. On the other hand, the genus 
Furtadoa of the subtribe Homalomeninae has a uniloculate ovary with basal pla- 
centation, but this genus differs from the genus Homalomena by the sterile pistil in 
the male portion of spadix, i.e., a male flower formed by a stamen and a sterile 
pistil. This interesting floral character is found in Homalomena mixta col- 
lected from the Malay Peninsula, the second species now assigned to the genus 
Furtadoa. 


Furtadoa mixtum (Ridley) M. Hotta, comb. nov. 

Homalomena mixta Ridley in Jour. Bot. 40: 36 (1902); Engler in Pflanzenr. ITV. 23 Da: 80 (1912); 
Furtado in Gard. Bull. Str. Settl. 10: 209 (1939). 
MALAYA. PAHANG: Tahan woods, 1891, Ridley s.n. (SING, holotype of H. mixta Ridley). 


This species has elliptic and wider leaf lamina than Furtadoa sumatrensis and 
seems to be a forest-floor aroid. 


Acknowledgments 


It is a pleasure to record my thanks to the many persons who helped me in my 
work. I am grateful to Dr. Kuswata Kartawinata of Herbarium Bogoriense and Dr. 
Chang Kiaw Lan of the Singapore Herbarium for the use of facilities under their 
care and for kindly helping with my herbarium work; Dr. Amsir Bakar, Head of 
Indonesian counterparts of SNS, Andalas University, and Dr. S. Kawamura, 
Leader of the SNS project, Kyoto University for their encouragement. Mr. Rusjdi 
Tamin and students of Andalas University kindly co-operated during my recurrent 
field work in West Sumatra. Dr. H. Okada has carefully made chromosome counts 
of the new species of Homalomena. 


References 


Engler, A. (1912). Araceae-Philodendroideae-Homalomeninae und Schismato- 
glottidinae. Engler’s Pflanzenr. IV. 23 Da: 1-82. 


54 Gard. Bull. Sing. 38(1) (1985) 


Furtado, C. X. (1939). Araceae Malesicae II: Notes on some Indo-Malaysian 
Homalomena species. Gard. Bull. Str. Settl. 10: 183-238. 


Hotta, M. (1967). Notes on Bornean plants, Il. Acta Phytotax. Geobot. 22: 153- 
162. 


(1984). Check list of Araceae in Sumatra. Hotta (ed.): Forest Ecology and 
Flora of G. Gadut, West Sumatra, 91-114. Kyoto Univ., Kyoto. 


Okada, H. (1984). Chromosome counts of some plants collected from W. Sumatra. 


Hotta (ed.): Forest Ecology and Flora of G. Gadut, West Sumatra, 89-90. Kyoto 
Univ., Kyoto. 


A New Account of the Genus Horsfieldia (Myristicaceae), Pt 2* 


W.J.J.O. de WILDE 
Rijksherbarium, Leiden, The Netherlands 


EFFECTIVE PUBLICATION DATE: 31 AUG. 1985 


Contents 


Enumeration and description-of the species 6-45 .....................2..0ceseccsecncceeceteceeceescnscescceess page 55 


6. Horsfieldia irya (Gaertn.) Warb. Fig. 1A(6); 2 II; 6 


Myristica irya Gaertn., Fruct. 1 (1788) 195, tab. 41; Hook.f. & Thoms., Fl. Ind. (1855) 159; A. DC., 
Prod. 14 (1856) 202 (excl. M. exaltata, p.p., see under Endocomia, Blumea 30 (1984) 173; King, Ann. 
Roy. Bot. Gard. Calc. 3 (1891) 308, pl. 141, 141-bis. — H. irya (Gaertn.) Warb., Mon. Myrist. (1897) 
317 (incl. vars. or forms ceylanica, javanica, malayana, wallichii, moluccana, siamensis), t. 22 fig. 1-4; 
Sinclair, Gard. Bull. Sing. 16 (1958) 382, fig. 33, Pl. IX-A; 28 (1975) 61; Back. & Bakh. van den 
Brink, Fl. Java 1 (1963) 138. — Type: Gaertner’s drawing. 


M. javanica B1., Bijdr. (1825) 576; Rumphia 1 (1835) 190, t. 62. — Type: authentic Blume’s specimens 
not found in L; tab. 62 (C fl., fr.). 


M. spherocarpa Wall., Pl. As. Rar. (1830) 79, t. 89. — M. irya var. wallichii King, Ann. Roy. Bot. 
Gard. Calc. 3 (1891) 309, pl. 141-bis, 3-5. — Type: Wallich Cat. 6796 (K-W). 


M. micrantha Wall., Cat. 6807 (1832), nom. nud. — Type: Wallich 6807 (K). 


M. lemanniana A. DC., Ann. Sci. Nat. Sér. 4, 4 (1855) 31, t. 4; Prod. 14, 1 (1856) 203. — H. lemanniana 
(DC.) Warb., Mon. Myrist. (1897) 326. — Type: Lemann s.n. (G, n.v.). 


M. subglobosa Migq., Fl. Ind. Bat. Suppl. 1 (1861) 383. — M. globularia Bl. var. subglobosa (Miq.) 
Miq., Ann. Mus. Lugd. Bat. 1 (1864) 206 — H. subglobosa (Miq.) Warb., Mon. Myrist. (1897) 328 
(for the original syntype only) — Type: Sumatra, Diepenhorst Hb. 2148 (U), Teysmann Hb. 3189 (VU). 


M. vrieseana Miq., Ann. Mus. Bot. Lugd. — Bat. 2 (1865) 49. — M. irya var. longifolia King, Ann. 
Roy. Bot. Gard. Calc. 3 (1891) 309, pl. 141-bis, 1-2. — Type: de Vriese s.n. (L). 


H. labillardieri Warb., Mon. Myrist. (1897) 283, t. 21 f. 1-2. — M. labillardieri (Warb.) Boerl., Hand. 
Fl. Ned. Ind. 3, 1 (1900) 85. — Type: Java, Hb. — Labillardiere s.n., male flowers (B, +; iso; FI, see 
note by Sinclair p. 89). 


H. acuminata Merr., Phil. J. Sc. 17, 1920 (1921) 253; En. Phil. Fl. Pl. 2 (1923) 181. — Type: de Mesa FB 
27507 (PNH, n.v., probably destroyed). 


H. nunu Kanehira, Trop. Woods 29, 5 (1932) nom. nud.; Bot. Mag. Tokyo 46 (1932) 451; Fl. Micr. 
(1933) fig. 32; Enum. Micron. Plants, in J. Dept. Kyushu Imp. Univ. 4, 6 (1935) 319. — Type: 
Kanehira 1303, 1304 (n.v.). 


H. amklaal Kanehira, Bot. Mag. Tokyo 47 (1933) 670; Fl. Micr. (1933) 109, fig. 31, pl. 16. — Type: 
_Kanehira 1944 (FU, n.v.), 1978 (FU, n.v.), 2058 (FU, n.v., iso: K & P), 2059 (FU, n.v.). 


H. congestiflora A.C. Smith, J. Arn. Arb. 22 (1941) 64. — Type: Brass 8010 (A, n.v.; iso: BM & L). 


Tree 10-25(-40). Twigs terete or often drying flattened towards apex, usually 
thinly ridged, (2-)3-10(-30) mm diam., glabrescent, tomentum minute to conspi- 


*Continued from Gdns’ Bull. Sing. 37(2): 179. 


55 


56 


Horsfieldia irya (Gaertn.) Warb. 

a, leafy twig apex, note whitish blotched leaves, x 1%; b, twig portion with male inflorescence, 
note ridged twig, X 42; c, mature male flower bud, lateral view, x 12; d, male flower, 
longitudinal section, showing androecium, X 25; e, androecium, longitudinal section, schema- 
tic, X 25; f, twig portion with female inflorescence, X 12; g, mature female flower, x 6; h, 
ditto, opened, showing glabrous ovary with minute 2-lipped stigma, X 12; i, twig portion with 
infructescence, note spherical fruits. — a & b from NGF 22319; c-e from Kostermans 24385; f-i 
from FRI 3044. 


New account of Horsfieldia 2 . SF 


cuous (New Guinea and Pacific Isls.), grey to rusty, of mixed dendroid hairs 
0.1-0.5(-1.0) mm; bark often coarsely striate, often + blackish, when older not 
flaking; lenticels usually conspicuous. Leaves in 2 rows, abaxially often + curved 
especially towards the tip, membranous, elliptic-oblong to lanceolate, 10-30(-35) 
x 3-7(-9) cm, base rounded to attenuate, tip acute-acuminate; upper surface 
drying dull greenish-brown to blackish-brown, usually finely pustulate with paler 
stipples and almost always with larger irregular whitish marks of unknown origin, 
lower surface early glabrescent to glabrous, without dark dots; midrib slender 
above, flattish; nerves 10-20 pairs, very thin and flattish above, inconspicuous, the 
marginal arches usually not distinct; tertiary venation forming a lax network, faint 
above, thin though distinct beneath; petiole 7-16 x 1.5-3(-4) mm; leaf bud c. 
10(-15) x 2-3 mm, pubescent with hairs c. 0.1-0.5(-1.0) mm. Inflorescences densely 
tomentose with hairs 0.1-0.5(-1.0) mm long, persistent or glabrescent, in GC: 3-4 
times ramified, many-flowered, c. 4-18 x 3-7(-10) cm; in Q: c. 2-6(-8) cm long, 
2(-3) times ramified; common peduncle 0.5-4.5 cm long; bracts acutish, 1.5-4 mm 
long, caducous. Flowers in CO in clusters of 3-10, in 9 usually solitary or a few 
together, perianth glabrous or at base glabrescent, perianth 2-valved; pedicel 
pubescent or glabrescent, at base not articulated. Male perianth subglobose or + 
transversely ellipsoid, somewhat laterally compressed or not, c. 1.0-1.3(-1.5) x 
(1.0-)1.2-1.5(-2.0, Indo-China) mm, apical part broadly rounded, at base rounded 
or short-tapering; pedicel slender, (0-)0.1-1(-1.5) mm, inarticulate at base; perianth 
at anthesis cleft to c. 42-7, valves c. 0.2 mm thick. Androecium broadly obovoid, + 
broadened transversely, c. 0.8-1.2 xX 1.0-1.5 mm; anthers 6-9(-10), Indo-China), 
not closely touching, c. 0.5-0.8(-1.0) mm long, towards apex incurved and free for 
c. 0.2-0.3 mm, dorsally attached to the broadly concave, often + saucer- or 
cup-shaped androphore c. 0.4-0.5 x (0.5-)0.6-1.0 mm, tapered towards the base. 
Female perianth obovoid or ellipsoid, c. 1.5-2.3 x 1.3-2.0 mm, at anthesis cleft to c. 
4-13, valves c. 0.3 mm thick, pedicel 1-4 mm long; ovary broadly obovoid, 
glabrous c. 1.2-1.5 X 1.0-1.3 mm, stigma minute, c. 0.05 xX 0.1 mm. Fruits 2-8 per 
infructescence, globose, 1.5-2.2 cm diam., glabrous, with the surface finely granu- 
lar, without larger tubercles or lenticels, drying dark brown to blackish, dry 
pericarp c. 1-2 mm thick; stalk 5-10 mm long, perianth not persisting. 


Distribution. From Ceylon through Malesia to the Solomon Isls.: Ceylon, Bur- 
ma, Andaman Isls., Nicobar Isl., S. Indo-China (Cochin-China), Cambodia, Thai- 
land, Malaya, Singapore, Sumatra, Java, Borneo, Celebes, Moluccas, C.(?) & S. 
Philippines, New Guinea, Caroline Isls., Solomon Isls.; no collections seen from 
the Lesser Sunda Isl. and N. Philippines. 


CEYLON: Davidse & Sumithraarachi 8119; Gardner (Hb. Hooker) 748; King’s Coll. (1884); Koster- 
mans 24385, 27202; Thwaites C.P. 221, 2620; Walker s.n.; Wall., Cat. 6804 (C’, part of the material of 
M. exaltata, not the lectotype); Waas 1272; Worthington 487, 517, 576, 686, 1848, 6351. 

ANDAMAN Isls.: Balakrishnan & Bhargava 3622; Kurz s.n.; Parkinson 1050. 

NICOBAR Isl.: Nair 3512. 


BURMA: Wallich 6804 C (Moulmein, C fls.; part of the original material of M. exaltata, not the 
lectotype). 


THAILAND: Kerr 4108 (A, B, C), 11419, 13883, 14252, 18599, 18907, 19036; Lakshnakara 615; 
Larsen c.s. FHB. 31253; Marcan 232, 720, 971, 1978; Maxwell 75-61; 75-1026; Put 623, 1580; Rabil 218, 
273; Smith 348; see further Sinclair’s list (1975, p. 62). 


CAMBODIA: Poilane Ch. 158. 


58 Gard. Bull. Sing. 38(1) (1985) 


VIETNAM (SOUTH): Miiller 1020; Pierre 5745; Poilane 769, Thorel 1186 (p.p., other specimens of 
the same number are the type of H. thorelii). 


MALAYA: Griffith 4357; Kep. FN 94710; Ridley 4957; Wallich 6807 — Kedah: Wyatt-Smith KFN 
71179 — Perak: FRI 3044; King’s coll. 7447; Ridley 3043; SFN. 33240 — Kelantan: Ridley 7206; Shah & 
Kadim MS 550 — Trengganu: SFN 40739. — Pahang: Evans s.n. — Selangor: SFN 34145. — Malacca: 
Kep. FN 94710; Maingay 1292, 2944, 3075. — Johore: Corner 25856, 25964, 28493; Maxwell 80-142; 
Ridley 897, 11328. — Penang (etc.): Curtis 936; Kep. FN 80999; Ridley s.n. 


SINGAPORE: Ridley 4814, 8957; SFN 339448, 40202; Sinclair s.n. (1953). 


SUMATRA: Ashton 15361; b.b. E. 868, 5208, 21463, 28460; Buwalda 6804; Forbes 3197; Grashoff 
1088; Horsfield s.n.; Praetorius s.n.; Rahmat si Toroes 3961 — Simeulué Isl.: Achmad 60, 362, 830, 1732 
— Mentawai Isls. (Sipora): [boet 487, Ridley 14760 — Riau: b.b. 20382. 


JAVA (W, C & E): Backer 27971; b.b. 1177, Boerlage s.n.; Buwalda 3001A, 3076 (= FRI. Ja. 4214); 
Coert 1489; Hoogerwerg a 1954; Junghuhn s.n., 33, 49, 875; Koorders 5211 fB, 5213 8, 5215 8, 5224 f, 
5265 f, 12274 B, 13493 B, 15521 8, 24784 B, 25392 B, 27479 B, 28353 B, 38882 8; Kostermans (Unesco) 52, 
19281; Soepadmo 279; Van Steenis 5274; Nengah Wirawan 429. 


BORNEO. Sarawak: Haviland & Hose 3305, 3305B; S. 16805, 18120, 18864, 34153, 36084 — Brunei: 
(Ashton) BRUN 5553 — Sabah: Amdjah 1, 94, 846; Elmer 20013, 21032; Rajuyap A 463; SAN. 31, 
18726, 21768, 30297, 34497, 47151, 75131, 84575, 90864 — W. Kalimantan: Hallier 1021, 1043; 
Teysmann 8676, 8680, 8683 — C. Kalimantan: Veldkamp 8256 — S. Kalimantan: Korthals s.n. —E. & 
SE. Kalimantan: b.b. 2114, 19042, 25129, 29415, 34244; Kostermans 4006, 5047, 21226, 21456. 


CELEBES: b.b. 22986; Forster (?) 329; Koorders 18157 8; Meijer 10139; Noerkas 305; Prawiroat- 
modjo & Soewoko 1777; Teysmann (11897). 


MOLUCCAS: Aupjé 33, 50; de Vogel 3807; de Vriese s.n., s.n. (87). 


PHILIPPINES. Palawan: Elmer 12682, 12684; Merrill 9208 — (Luzon, Mindanao): Cenabre FB 
29146, 29176; Cruz FB 23887; Natividad FB 25751; (Ramos & Edano) BS 36770, 41198; Vidal 3567; 
Wenzel 3023. 


NEW GUINEA. Irian Jaya (West New Guinea): Zippel s.n.; b.b. 22530, (Kostermans 127) 33354; 
BW 393, 873, 3288, 4436, 5165, 5348, 5838, 5840, (Schram BW) 6082 (in K), 6563, 6595, 7258, 10827, 
11870; van Royen 4675, 5101 — Papua New Guinea: Brass 8010; Craven & Schodde 776; Hartley (TGH) 
9738; Hollrung 657 (in P, not in K); Hoogland 4213, 4650; LAE 74298; NGF 16320, 22319, 25577, 
35309, 37574, 47300. 


CAROLINE Isls.: Kanehira 1303, 2058; Masahiko Takamatsu 392; St. John 21446; Stone 1901. 


SOLOMON Isls.: BSIP 39, 909, 910, 1663, 1664, 2014, 3668, 4743, 5286, 5788, 5855, 6037, 6119, 
6948, 10614, 13211, 15441, 15806, 17057, 17275; Comins 355; Kajewski s.n., 2444. 


Ecology. A tall tree in primary and (old) secondary forest. Most frequent coastal 
or riverine, on alluvial (sandy, loamy, or clayey) soils, but also found more inland; 
0-450 m. Mentioned mostly from wet or marshy localities at low altitudes, e.g., 
coastal swamp forest, sandy alluvium behind the beach, open, low marshy ground 
(with sago-palms), swamp forest, river banks below tidal limits, seasonal swamps, 
periodically inundated forest, coastal shrubberies (on limestone), river flood-plains 
with mud soil, etc., but also recorded from undulating country, and well-drained 
soils (e.g., in Ceylon and Solomon Isls.). From Ceylon recorded from the “dry 
zone’. In Thailand in riverine evergreen forest. Flowers and fruits apparently 
throughout the year, but this is likely correlated with regional and local climatic . 
conditions. 


Vernacular names. Amklaal, Nunu (Palau Isls.); Aininiu, Ainynu (Kwara’ae 
lang., Solomon Isls.); more vernacular names will be published in Flora Malesiana. 


New account of Horsfieldia 2 59 


Uses. Rare in record that fruits are edible; fruits recorded as eaten by monkeys in 
Ceylon. 


NOTES 


1. Fieldnotes. A tall tree with straight bole; crown described as with several big 
limbs each monopodially branched, or narrow and with slender drooping branches 
near the top; bole often recorded as fluted, or with prop roots, or, usually, with 
buttresses up to 3 m high, 2 m out, and up to c. 10 cm thick, but also recorded as 
without buttresses. Bark sometimes noted as smooth, often as fissured or cracked, 
or mostly as flaking or peeling off in small pieces; inner bark cream or whitish, to c. 
7 mm thick; sapwood cream or whitish, ochreish, straw-coloured, or pinkish; 
heartwood absent or only slightly darker, pale brown; wood rather soft. Flowers 
yellow, dark yellow, or orange-yellow, once recorded as reddish; strongly sweet- 
scented, once recorded as unscented. Fruit yellowish, greenish orange, to orange- 
red, or red; the aril usually orange or orange-red, rarely recorded as red. Fresh fruit 
recorded as large, up to c. 2*%4 cm diam. The seed is reported to contain an air 
chamber, facilitating dispersal by floating. 


2. Variability. H. irya is a homogeneous species, well characterized by its small 
subglobose male flowers of c. 1 mm diam. (in Indo-China up to 1.6 mm diam.), 
with typical broad and deeply concave androecium with tapered, distinct (i-e., 
relatively large) androphore. The fruits (and seeds) are perfectly spherical, glab- 
rous (ovary glabrous). The twigs are usually thinly ridged from petiole to petiole. 
Very characteristic are irregular whitish marks of unknown origin, almost always 
present on the older leaves. The leaves have a lax reticulation. Variation abounds 
in the tomentum: short-haired; sometimes seemingly glabrous specimens are pre- 
dominant in Ceylon, SE. Asia, and W. Malesia and the Moluccas; in New Guinea 
and the Solomon Isls. most specimens have a conspicuous, often wooly, tomentum 
of hairs to c. 1 mm long on the twig apex, leaf bud, and inflorescences. 


7. Horsfieldia spicata (Roxb.) Sinclair Fig. 1A(7); 7 


Myristica spicata Roxb., Fl. Ind. 3 (1832) 847; (ed. 1874) 744; Warb., Mon. Myrist. (1897) 271 in obs. 
sub H. smithii Warb. — H. spicata (Roxb.) Sinclair var. spicata, Gard. Bull. Sing. 28 (1975) 112, 113, 
p.p. — Type: Roxburgh’s description. 


M. canariformis B\., Rumphia 1 (1837) 190 — H. canariformis (Bl.) Merr., Interpret. Rumph. (1917) 
230 — Based on: Palala quarta, P. canariformis, P. dentaria Rumph., Hb. Amb. 2, 10 (1741) 27 t. 8; 
see Sinclair, 1975, 162-165. 


H. batjanica Warb., Mon. Myrist. (1897) 275, tab. 21, 1-4 — M. batjanica (Warb.) Boerl., Hand. FI. 
Ned. Ind. 3, 1 (1900) 85 — Type: Introduced by Teysmann in hortus Bogor. (B, lost; iso: FI, n.v.; 
original tree still cultivated in Bogor, and collected sub Kostermans 11186, Rastini (220), Sinclair 
10035). 


H. roxburghii Warb., Mon. Myrist. (1897) 277, tab. 21, 1-2. — M. roxburgii (Warb.) Boerl., Handl. Fl. 
Ned. Ind. 3, 1 (1900) 85 — Type: Smith in Hb. Roxburgh (BM; B, BR, n.v.; Q fl., orig. Ternate); 
Culta in hortus Bog. (C fl., orig. Ambon) (B, FI, n.v.; lecto: P, tree still in cultivation in Bogor sub 
no. IV. G. 90, collected under Sinclair 10037). 


H. parviflora auct. non (Roxb.) Sinclair: Sinclair, Gard. Bull. Sing. 28 (1975) 82, p.p. 


Tree 2.5-20 m. Twigs terete, not ridged, towards the apex 2-4 mm diam., 
tomentum greyish, with hairs c. 0.1 mm or less, early glabrescent; bark striate, pale 
brown to whitish brown, usually contrasting with the blackish colour of dried 


Fig. 7. 


Horsfieldia spicata (Roxb.) Sinclair 

a, leafy twig with male inflorescences, x 12; b, mature male flower, lateral view, X 6; c, male 
flower, opened, showing androecium, x 6; d, male perianth, inner side, showing impression of 
androecium, X 6; e, androecium, longitudinal section, schematic, x 6; f, twig portion with 
female inflorescence, X 12; g, mature female flower, lateral view, < 6; h, ditto, opened, 
showing glabrous ovary and 2-lipped stigma, 6; i, twig portion with infructescence, x 2. — 
a-d from Beguin 1407; e from Teysmann s.n. (Ambon); f-h from Kostermans s.n. (Hort. Bog. 
sub IV.H. 13); i from de Vogel 3206. 


New account of Horsfieldia 2 61 


petioles and inflorescences, when older not flaking, usually with few coarse len- 
ticels. Leaves in 2 rows, membranous, elliptic-oblong to oblong, 8-30 x 2.5-10 cm, 
base attenuate, top acute-acuminate, often densely speckled by paler irregular 
pustules of unknown origin, especially beneath; upper surface drying dull greenish 
brown to brown, lower surface with very minute scales as on the leaf bud, very 
early glabrescent, i.e., glabrous; without larger blackish dots, but often with dense, 
very minute, blackish dots; midrib flattish above; nerves 11-17 pairs, thin and 
flattish above, tertiary veins very thin, distinct or not on both surfaces; petiole 
10-20 x 1.5-3 mm, drying blackish; leaf bud slender, c. 10 xX 1.5-2 mm, densely 
pubescent with grey-brown hairs c. 0.1 mm long or less. Inflorescences drying 
blackish, usually slender, spike-like, early glabrescent, not ramified or the lateral 
branches only up to c. 2(-5) mm, common peduncle c. 1-3 cm, not many-flowered, 
in CO: 4-10 X c. 1 cm, in Q: c. 2-3 cm long; bracts bluntish, 0.5-1 mm, caducous. 
Flowers solitary or 2 together in 9, up to 3 together in ©’, very early glabrescent; 
perianths 2-valved; pedicel glabrous (glabrescent), at base inarticulate. Male 
perianth + obovoid or short-pear shaped, in lateral view the upper part subcircular 
to reniform, laterally rather compressed, about as long as broad or slightly broader 
than long, 2.3-3 x 3-3.5 mm, the upper part broadly rounded, base + tapering into 
the much tapered pedicel 1.5-3 mm long; perianth at anthesis split to or nearly to 
the base, valves c. 0.2 mm thick. Androecium reniform, rather well compressed, 
upper part broadly rounded, at base broadly attached, 1.5-1.8 x 2.5 mm, 
androphore + absent; anthers (12-)16-22, closely set, + septate only in young state, 
free apices 0.1-0.4 mm, little to strongly incurved into the apical cavity which 
reaches to c. %4-'4(-%4, see notes). Female perianth subglobose, 2-2.5 x 2.2-2.8 
mm, at base passing into the somewhat tapered pedicel 1.5-2.5 mm, at anthesis split 
to c. 2-%, valves 0.4-0.5 mm thick; ovary broadly ovoid, glabrous, c. 1.5 x 1.5 
mm, stigma minute, faintly 2-lobed, c. 0.1 mm high. Fruits 1-5 per infructescence, 
short-ellipsoid, apex and base rounded, 1.5-2.0 xX 1.2-1.8 cm, glabrous, drying 
blackish, without pustules, pericarp c. 1.5 mm thick; stalk 4-10 mm long; perianth 
not persisting. 


Distribution. Moluccas: Morotai, Halmahera, Ternate, Bacan Isl., Ambon. 


Cultivated (origin Bacan Isl., Ambon): Kostermans 11186; Rastini (220); Sinclair 10035, 10037; Hb. 
Hasskarl s.n. (Teysmann s.n., 1868, orig. “E. Java’’). 


MOROTAI: Kostermans 1014, 1157, 1218, 1256; Lam 3499. 

HALMAHERA: Pleyte 409, p.p.; de Vogel 3206, 3289, 3334, 3367, 3434, 3490, 3496, 4391, 4467. 
TERNATE: Beguin 1407. 

BACAN: b.b. 32787; Hb. Hance s.n.; de Vogel 3529, 3531, 3748. 


AMBON: Buwalda 6141; Robinson 240, 1885; Teysmann s.n. (1867). 


Ecology. Alluvial soils, deep clay, soil rich in humus, and porous volcanic soil 
over schists; 0-1000 m. Flowers and fruits throughout the year. 


Vernacular names. Anunu magilioro (Halmahera), Onguaka (Tobaro lang., 
Halmahera). 


Uses. According to the label of de Vogel 3206, the outer bark, mixed with 
“Kuleman” (a different species of Horsfieldia), is used for curing hepatitis. 


Fig. 8. 


62 


Horsfieldia inflexa de Wilde 

a, twig portion with male inflorescences, note lined twig, X 12; b, mature male flower, opened, 
showing androecium, X 12; c, androecium, longitudinal section, schematic, x 12; d, twig 
portion with female inflorescences axillary to fallen leaves, x 1%; e, mature female flower, 
lateral view, X 12; f, ditto, opened, showing glabrous ovary with broad-lipped stigma, x 12; g, 
twig portion with infructescences, X 12; h, portion of lower leaf surface with scattered 
dark-coloured non-traumatic cork warts as blackish dots, X 12. — a-c from LAE 52866, type; 
d-f from van Royen 3166; g & h from LAE 52862. 


:/ he 


New account of Horsfieldia 2 63 


NOTES 


1. Fieldnotes. Flowers greenish-yellow or ochreish yellow. Mature fruit orange; 
aril bright red. Bark often recorded as unfissured, peeling off. Exudate watery, not 
or only slightly reddish coloured. Sapwood usually creamish, gradually passing into 
the darker coloured heartwood. Buttresses recorded as present and up to 50 cm out 
and high, or as absent. 


2. This species is clearly recognizable at first glance by its spike-like infloresc- 
ences drying blackish, and by the pale twigs rather contrasting with the blackish 
colour of the dried petioles and the inflorescences. As presently circumscribed by 
me this species is used in a much narrower sense than accepted by Sinclair, who 
included also specimens from Celebes, Philippines, Lesser Sunda Isls., and Banda 
and Aru Isls., now referred by me to various different species. 


3. Possibly its most closely related species is H. moluccana. Apart from the 
characters as used in the key, and pointed out in note 2, H. spicata differs from H. 
moluccana by its more membranous leaves. See also note 4. 


4. Intermediate specimen. Teysmann s.n. (in L), from Ambon, is intermediate 
between H. spicata and the related H. moluccana. It has the diagnostic conspi- 
cuously pale twigs, contrasting with the blackish petioles and inflorescences, but 
these latter structures are rather ramified, not spicate, with ramifications at the 
base c. 6 mm long. The androecium of the flowers of this specimen are hollow for 
c. ¥4 and the anthers at one side of the androecium curve into this apical cavity. 
Possibly the specimen is a hybrid. 


5. Sinclair (1975, p. 122) discussed elaborately the typification of Roxburgh’s 
name which is accepted here. 


8. Horsfieldia inflexa de Wilde, sp. nov. Fig. 1A(8); 8 


Horsfieldia species perianthiis masculis usque ad basin 2-valvibus atque antheris inflexis, ex affinitate 
gregis H. sepikensem et H. moluccanam amplectens, sed differt virgis valde porcatis atque foliis subtus 
punctatis. — Typus: Streimann & Martin LAE 52866 (L). 


Tree 10-21 m. Twigs distinctly ridged from petiole to petiole and distinctly 
angular especially in the apical portion, 2-5(-7) mm diam., early glabrescent, 
tomentum grey-brown, of hairs up to 0.1 mm; bark indistinctly striate, when older 
not flaking, lenticels abundantly present but usually not very conspicuous. Leaves 
in 2 rows, thinly chartaceous, elliptic-oblong to oblong, broadest at about or + 
above the middle, 8-20 x 2.5-7.5 cm, base attenuate, tip bluntish to acute- 
acuminate; upper surface drying olivaceous to blackish brown, not or indistinctly 
pale-pustulate, lower surface early glabrescent and with + regularly scattered dark 
brownish dots (lens X 10!); midrib above slender, flattish or slightly raised; nerves 
8-13 pairs, above thin and flat, indistinct, beneath with the marginal arches fairly 
regular but indistinct; tertiary venation forming a rather lax network indistinct on 
both surfaces; petiole 14-30 x 1.5-2.5 mm; leaf bud 10-14 x 1.5-2 mm, with 
tomentum, hairs c. 0.1 mm long or less. Inflorescences glabrescent or with scat- 
tered minute scale-like hairs less than 0.1 mm; in CG: 2(-3) times ramified, the 
primary branches rather spike-like, c. 3-10 x 1.5-4.5 cm, in 9: 3-5 cm long; 
common peduncle 1-12 mm long; bracts elliptic, 1-3 mm long, with fimbriate 
margin, caducous. Flowers solitary or up to 4 together, glabrous; perianth 2-valved; 
pedicel glabrous, at base inarticulated. Male perianth subglobose, generally slightly 


64 Gard. Bull. Sing. 38(1) (1985) 


broader than long, not or but little laterally compressed, sometimes (together with 
the pedicel) slightly pear-shaped, c. 2.2-2.5 x 2.5-3 mm, upper part broadly 
rounded, lower part rounded or slightly tapering; pedicel 1-2 mm long, glabrous; 
perianth at anthesis split to almost reaching the base (c. 10), valves 0.2-0.5 mm 
thick, not collapsing on drying. Androecium bluntly quadrangular in outline, 
sometimes broader than long, broadly rounded above, laterally compressed, c. 1.5 
x 1.5-2.0 mm; anthers 10-12(-14?), septate when young, c. 1.5-2 mm long, free 
apices c. 0.7-1.0 mm (i.e., anthers upper half free), at one side of the androecium 
strongly incurved into the hollow almost reaching the base; androphore 0-0.1 mm 
long. Female perianths ovoid-ellipsoid, c. 2.0-2.5 x 2 mm, cleft at anthesis to c. %, 
valves c. 0.5 mm thick; pedicel 1-1.5 mm long; ovary ovoid, c. 1.5 x 1.2 mm, 
glabrous, stigma sessile, minutely 2-lobed, c. 0.2 mm broad. Fruits solitary or up to 
6 per infructescence, ellipsoid, top rounded to subacute, base + rounded, c. 2.0 x 
1.4-1.5 cm, glabrous, drying dark brown, without or with small tubercles or 
lenticels only; dry valves c. 1.5 mm thick; stalk 2-6 mm long; perianth not per- 
sisting. 


Distribution. Northern part of New Guinea. Irian Jaya: Vogelkop, Geelvink 
Bay, Jayapura; Northern Papua New Guinea: West Sepik. 


NEW GUINEA. Irian Jaya (northern) (incl. Vogelkop, Japen Isl., Meos Noem): Aet & Idjan (exp. 
van Dijk) 389; b.b. 21812, 25724, 30376, 30617, 30945; BW (Schram) 6046, (Iwanggin) 10021, (Moll) 
11672; Pleyte 736; van Royen 3166. — Papua New Guinea (northern), West Sepik: (Streimann & 
Martin) LAE 52862; 52866. 


Ecology. Primary and old secondary forest on alluvial soils, e.g., sandy clay, also 
in hilly forest, swamp forest; 0-400 m alt. Flowers throughout the year, fruits from 
September to November. 


Vernacular names. Kamopi (Roberbai, Japen Isl.), Madak (Mooi lang., Vogel- 
kop), Teenjak (Tehid lang., Vogelkop). 


NOTES 


1. Fieldnotes. Slender tree, buttresses absent. Bark shallowly longitudinally 
fissured, not or slightly peeling off; slash reddish brown, sapwood whitish red or 
cream, heartwood not differentiated. Flowers green, turning yellow, fragrant. 
Fruits yellowish-orange. 


2. Besides a few species like H. glabra and H. punctatifolia from Western Malesia 
this is the only other Horsfieldia but the only one in New Guinea with specifically 
typical largish blackish dots on the lower leaf surface. These dots are apparently of 
the same nature as those in series Punctatae of the genus Knema, and have been 
identified as cork warts of non-traumatic origin. 


3. Specimens of the present new species were included by Sinclair in his H. 
spicata var. sepikensis (Mkgf.) Sinclair, a taxon which is presently regarded as 
representing several distinct species, among which H. sepikensis, H. moluccana 
(var. petiolaris) and H. olens. 


4. H. inflexa is obviously closely allied with H. moluccana, and can also be 
confused with the closely related species H. angularis and H. basifissa. 


New account of Horsfieldia 2 65 


H. moluccana has generally rather pear-shaped male flowers (incl. pedicel), which 
may also be the case in certain specimens of our present H. inflexa, e.g., BW 6046 
from Vogelkop; H. moluccana differs, however, by its terete or only faintly lined 
twigs, and its non-punctate leaves. H. inflexa may resemble H. moluccana very 
much in the general shape and texture of the leaves, including the relatively long 
petioles. H. angularis differs by its non-punctate leaves, hairy flowers, the 
androecium with a central narrow crevice and its straight anthers. H. basifissa has, 
in contrast, terete or only faintly ridged twigs, differing further by various charac- 
ters of the male flower including the androecium. 


5. The specrments BW 10021 and b.b. 30617 from Japen Isl. resemble H. 
parviflora in their rather unbranched spike-like inflorescences; H. parviflora has 
more pronounced pear-shaped flowers and twigs which are paler and not angular. 
In most of the specimens of H. inflexa, the male inflorescences are distinctly 
ramified whereas the lateral branches are almost unbranched and are spike-like. 


9. Horsfieldia moluccana de Wilde, sp. nov. Fig. 1A(9) 


Horsfieldia olivaeformis Warb., Mon. Myrist. (1897) 352, t. 23 fig. 1-2; Markgraf, Bot. Jahrb. 67, 2 
(1935) 152, p.p. — Myristica olivaeformis (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 87 — 
Type: Irian Jaya, Sorong (Vogelkop), Beccari 17] (FI, n.v., identity not sure, see notes). 


Horsfieldia species perianthiis masculis 2-valvibus pyriformibus atque antheris inflexis, eis H. 
spicatae similibus, ab eo differt virgis in sicco dense brunnescentibus atque inflorescentiis valde ramosis 
non spicatis. — Typus: Kostermans 673 a (L). 


Tree, 8-20(-30) m. Twigs terete, not ridged, or sometimes slightly angular, 
towards the apex 2-5 mm diam., early glabrescent; tomentum with hairs 0.1-0.3 
mm; bark striate, when older not flaking, lenticels conspicuous to inconspicuous. 
Leaves in 2 rows, thinly chartaceous, elliptic-oblong to oblong, (6-)8-25 x 
2.5-8.5 cm, base attenuate, tip acute-acuminate; upper surface drying olivaceous 
to brown, usually minutely pale-pustulate, lower surface early glabrescent, without 
dark brown dots; midrib slender above, flat; nerves 6-15 pairs, above thin, flat, 
indistinct, beneath with the marginal arches indistinct; tertiary venation + fine, 
indistinct on both surfaces; petiole 10-26 x 1-2 mm; leaf bud 6-12 x 1-2 mm, with 
hairs 0.1-0.3 mm. Inflorescences sparsely pubescent, hairs stellate, c. 0.1 mm or 
less, in CG: (1-)2-3 times ramified (sometimes not or hardly ramified, i.e., spike- 
like, see notes), 5-11 x 2-5 cm, in 9 up to 5 cm long; common peduncle 5-20 mm; 
bracts + oblong, 1.5-4 mm long, thinly pubescent, caducous. Flowers from solitary 
to 4 together, glabrous; perianths 2-valved; pedicels glabrous, inarticulate at base. 
Male perianth (incl. pedicel) pear-shaped, laterally much to little compressed, 
about as broad as long to slightly broader than long, 1.5-2.5(-3) « 2.2-3.8 mm, 
upper part broadly rounded, the lower 3 tapering into the tapered pedicel 2-3(- 
3.5) mm; perianth at anthesis split to c. 74-5, valves 0.2-0.3 mm thick. Androecium 
laterally compressed, broadly transversely ellipsoid or kidney-shaped in outline, 
broadly rounded above, c. 1.1-1.5  1.4-2.8 mm; anthers (7-) 10-18, not septate, c. 
1.5-2.0 mm long, free apices 0.1-0.5 mm, only at one side of the androecium 
strongly incurved; androecium hollow for at least 73; androphore 0-0.1 mm long. 
Female perianths broadly ovoid-ellipsoid, 1.8-2.2 x 2.0-2.2 mm, cleft at anthesis to 
Y2-¥5, valves c. 0.3 mm thick; pedicel 2-2.5 mm long; ovary ovoid, glabrous, c. 1-1.5 
x 1 mm, stigma sessile, minutely 2-lobed, c. 0.1 mm high. Fruits solitary or 2-6 per 
infructescence, ellipsoid, top rounded to subacute, 1.3-2.8 x 1.1-1.7 cm, glabrous, 
drying brown or blackish, without or with sparse tubercles; dry valves 1-2 mm 
thick; stalk 2-5 mm long; perianth not persisting. 


66 Gard. Bull. Sing. 38(1) (1985) 


Distribution. Northern Moluccas, West New Guinea. 


NOTES 


1. A variable species with 4 varieties, closely related to H. spicata and H. 
tuberculata. With H. spicata there occur a few specimens intermediate to var. 
moluccana discussed in the notes. H. spicata has the generally deeply asymmetrical- 
ly incurved anthers in common with H. moluccana, but the former differs in the 
pale twigs and the (almost) spike-like male inflorescences. H. tuberculata has 
largely a solid staminal column, hollowed at the apex only for the upper 5-4. See 
further notes under the varieties. 


2. Unfortunately I have not seen the type of H. olivaeformis. If it turns out to be 
identical with the present new species, it would then have priority. Sinclair lumped 
this name in his large concept of H. spicata, from which the present new species is 
segregated. 


The type of H. olivaeformis, Beccari 171, has been described as having a glabrous 
ovary, the fruits rather narrow, c. 2.3 cm long and its pericarp thin, the leaf blades 
10-15 cm and petioles almost 10-15 cm long. In fruit size it agrees with var. robusta, 
known from the same area, but the petioles in that are distinctly longer than those 
described for H. olivaeformis. 


KEY TO THE VARIETIES 


la. Hairs of leaf bud rather woolly-rust pubescent, hairs 0.2-0.3 mm long. Vogelkop: Fak-Fak 
Sim H Paphagin dy bint «ed amin guia dp a hile s Wbediad ee ao bpeatebae vaerad REECE idk etcetera d. var. pubescens 


b.’ Hairs of leaf bud up-toc. 0. Bina oi as 02S 500, con apne gore toned eae er eae oe 2 
2a. Petioles 10-15 mm long. Male perianth 2.5-3 mm wide. Morotai, Obi Isls. .......... a. var. moluccana 
b. Petioles (10-) 15-25 mm long, generally longer in proportion to the smaller blade. New Guinea .... 3 


3a. Leaf blades 7-15 cm long. Fruits 1.3-1.8 cm long. Male perianth 2-2.5 mm wide ..... b. var. petiolaris 


b. Leaf blades 13-23 cm long. Fruits 2.2-3.0 cm long. Male perianth 3.5-3.8 mm wide ... ¢. var. robusta 


a. var. moluccana Fig. 1A(9) 


Leaf blades 9-22 x 4-8 cm; petioles (8-)10-20 mm. Tomentum of leaf bud 
composed of hairs c. 0.1 mm long or less. Male perianths 2.0-3.0 x 2.7-3.3 mm, 
pedicel 2-3 mm long. Fruits c. 1.5 cm long. 


Distribution. Northern Moluccas: Morotai, Obi Isls. 


MOLUCCAS (Northern). Morotai: Tankilisan (exp. Kostermans) 250 (= b.b. 33920); Kostermans 
673a, 899, 1513, 7888; Lam 3459, 3510. — Obi Isls.: de Vogel 4099, 4105, 4111, 4121, 4137, 4193, 4240, 
4253, 4298, 4303. 


Ecology. Well-drained forests on clayey soil, volcanic soil, alluvial soil rich in 
humus, also flat land just behind the mangrove; recorded from over limestone, or 
at base of serpentine-rock, or on very porous nickel-containing soil; 0-600 m alt. 
Flowers and fruits throughout the year. 


New account of Horsfieldia 2 67 


Vernacular names. Gosora (Ternate lang.), Kuleman (Morotai), Pala hutan 
(Malay lang.). 


NOTES 


1. Fieldnotes. Recorded as a straight tall tree, to 30 m. Bark peeling off or not. 
Once reported to have prop roots up to 1.5 m. Exudate from bark watery, turning 
pink, later turning brownish. Flowers yellow, once recorded as red. 


2. Sinclair included a large part of the present taxon in his concept of H. 
parviflora. In the present revision H. parviflora is accepted in a narrower sense, 
and is mainly characterised by more roundish (not pear-shaped) male perianths, 
and by shorter petioles. H. spicata is closely related but differs in the paler colour of 
the dried twigs, the generally spike-like male inflorescences and the more membra- 
nous leaves. See also note 3. 


3. Specimens intermediate to H. spicata. The male inflorescences of de Vogel 
4253, from Obi Isls at sea level, are spike-like. In all other aspects, including 
chartaceous leaves, short and robust male perianths and brown twigs, it is identical 
with other material from there. De Vogel 4193, also from the same source, has the 
inflorescences almost spike-like and are thus reminiscent of those in H. spicata. Its 
leaves are membranous and it may be a hybrid; it was collected from a secondary 
regrowth at c. 200 m. alt. One should note that the lectotype of H. roxburghii 
Warb. (in this treatment, a synonym of H. spicata) is from a tree cultivated in 
Bogor Botanic Gardens, with Ambon as its provenance; it has inflorescences which 
are rather branched, not strictly spike-like and thus it looks intermediate between 
H. moluccana and H. spicata. 


b. var. petiolaris de Wilde, var. nov. 


Gemmae indumentum brevissimum, maxime c. 0.1 mm longum, petiolo proportione longo, 1-2 cm 
longo, foliorum pagina c. 7-15 cm longa, perianthiis masculis 2-2.5 mm latis, fructibus c. 1.5 cm longis. 
— Typus: van Royen 5388 (L). 


Leaf blades 6-15(-19) x 2.5-6.5(-7) cm; petioles proportionally long, 10-20 mm. 
Tomentum of leaf bud composed of hairs up to 0.1 mm long. Male perianth 1.5-2.2 
x 2.2-2.4 mm and pedicel 2-3.5 mm long. Fruits 1.5-1.8 cm long. 


Distribution. Irian Jaya: Vogelkop; Islands in Geelvink Bay (Noemfoer, Meos 
Waar, Japen Isl.); Waigeo Isl. 


IRIAN JAYA: b.b. 30587, 32987; (Koster) BW 1018, 1201, 1283; BW 1295; (Kalkman) BW 6255; 
(Koster) BW 13535; van Royen 5388, 5396. 


Ecology. Locally common in forests on sandy or stony-clayey soils; Calophyllum- 
Ficus forest; 0-100 m. alt. Flowers and fruits throughout the year. 


Vernacular names. Beterohooi (Manikiong lang.), Kamojer (Noemfoer lang.), 
Mbowak (Tehid lang.), Sebohongwa (Manikiong lang. ). 


Uses. Fruits once reported as edible and sour. 


68 Gard. Bull. Sing. 38(1) (1985) 


NOTES 


1. Fieldnotes. Bark flaking. Flowers greenish. Fruits yellow or orange-yellow, 
seed aril red. 


2. Sinclair included the specimens of var. petiolaris in his H. spicata var. 
sepikensis (Mkgf.) Sinclair; here I have, however, kept H. sepikensis as a separate 
species, characterized by a 3-valved perianth. 


c. var. robusta de Wilde, var. nov. 


Gemmae indumentum c. 0.1 mm longum, petiolis c. 1.5-2.5 cm longis, foliorum pagina 13-23 cm 
longa, perianthiis masculus c. 3.5 mm latis, fructibus c. 2.5 cm longis. — Typus: van Royen & Sleumer 
6682 (L). 


Leaf blades 12-22(-25) x 3.5-8(-9) cm; petioles 12-26 mm. Tomentum of leaf bud 
consisting of hairs c. 0.1 mm long. Male perianths 2.5-3 xX 3.5-3.8 mm, pedicel c. 3 
mm. Fruits 2.2-3.0 cm long. 


Distribution. Irian Jaya: Vogelkop Penins.; Batanta Isl. 


IRIAN JAYA: (Moll) BW 9775; van Royen 3548; van Royen & Sleumer 6682. 


Ecology. Secondary and coastal forest, on limestone; 0-15 m. Flowers and fruits 
throughout the year. 


Vernacular name. Kamore (Biak dial.). 
NOTES 
1. Fieldnotes. Bark flaking. Flowers yellow, fragrant. Fruits yellow. 


2. Sinclair included the present variety in his H. spicata var. spicata. He has 
annotated on the type sheet (van Royen and Sleumer 6682) that it looked to him 
intermediate to H. sepikensis (in Sinclair’s sense). 


3. Var. robusta is a form similar to var. petiolaris but is coarser in all aspects: the 
leaves, flowers and fruits are all larger. It superficially resembles H. tuberculata, 
which differs in the generally shorter petioles, the androecium — the anthers not 
strongly inflexed into the cavity — as is the case in the present var. robusta. 


d. var. pubescens de Wilde, var. nov. 


Gemma lanata, e pilis c. 0.3 mm longis composita. — Typus: Vink BW 15370 (L; iso: K). 


Leaf blades (6-)8-14 x (2.5-)3-5 cm; petioles (9-)11-18 mm long. Tomentum of 
leaf bud + woolly, composed of hairs 0.2-0.3 mm long. Flowers not seen. Fruits c. 
1.3 cm long. 


Distribution. Irian Jaya: Vogelkop Penins. 


IRIAN JAYA. Vogelkop Penins.: BW (Iwanggin) 5640, (Schram) 6153, (Iwanggin) 10153, (Vink) 
15370. | 


New account of Horsfieldia 2 69 


Ecology. Common in primary and secondary forest on clayey soil or sandy clay 
over limestone; 50-300 m. alt. Fruits in March and May. 


Vernacular names. Kamorei (Biak lang.), Medak (Mooi lang.), Sémies, Simies 
(Maibrat lang.). 


NOTES 


1. Fieldnotes. Tree to 16 m; buttresses up to 1 m high, 0.5 m wide; bark strongly 
peeling. Inner bark reported as with much red and clear exudate. Wood white. 
Fruits light green. Most collections are from limestone. 


2. This variety appears to be almost identical with the var. petiolaris, except for 
the more woolly tomentum. In Horsfieldia, usually the nature of the tomentum has 
appeared to be of taxonomic significance. Flowers are not known. 


3. The bark of the older wood in the Kew duplicate of BW 15370 is one that 
flakes strongly; that in the Leiden specimen and some other collections, of older 


twigs behind the leaves, is one that does not or flakes minimally. 


4. Sinclair included the specimens of the present variety in his H. spicata var. 
sepikensis (Mkgf.) Sinclair, as those of the preceding variety. 


10. Horsfieldia parviflora (Roxb.) Sinclair Fig. 1A(10) 


Myristica microcarpa Willd. in Roem. & Usteri, Bot. Mag. 3, 9 (1790) 27; Sp. Pl. 4, 4, 2 (1806) 871 (excl. 
var. 8 = 7. H. spicata, incl. var. Y ). — Based on Palala ‘‘kitjil”, P. minima Rumph.; var. Y based on 
P. globularia Rumph.; identity doubtful, see Sinclair 1975, p. 168-170. 


M. parviflora Roxb., Fl. Ind. 3 (1832) 847; (ed. 1874) 744; Icones 2574. — H. parviflora (Roxb.) 
Sinclair, Gard. Bull. Sing. 28 (1975) 82. — Type: Roxburgh’s description and figure. 


M. tingens Bl., Rumph. 1 (1837) 190 — Horsfieldia sp. Merr., Int. Rumph. (1917) 231 — Based on 
Palala minima, P. tertia, P. tingens Rumph., Herb. Amb. 2, 10 (1741) 27, t. 7 f. A-B; see Sinclair 
1975, p. 161. 


M. globularia B\., Rumphia 1 (1837) 191, t. 64, fig. 2 (non Lamk.) — Pyrrhosa globularia (B1.) Hassk.. 
Cat. Pl. Hort. Bog. (1844) 174 — H. globularia (B|.) Warb., Mon. Myrist. (1897) 288, t. 21 (1-4). — 
Palala globularia (= P. quinta) Rumph., Herb. Amb. 2, 10 (1741) 28, t. 9 f. a-b (see Sinclair 1975, p. 
165-167). — Type: Blume’s figure, and Zippel s.n. (Ambon, ‘“‘mas’’), a sterile specimen. 


M. bivalvis Hook.f., Fl. Brit. Ind. 5 (1886) 107; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 307, pl. 139. 
— H. bivalvis (Hook.f.) Merr., Phil. J. Sc. Bot. 2 (1916) (issued Jan. 1917) 271; Sinclair, Gard. Bull. 
Sing. 16 (1958) 379, fig. 32, pl. VIII B. — Type: Murton 149 (K). 


H. globularia var. minahassae Warb., Mon. Myrist. (1897) 617. — H. minahassae (Warb.) Koord., FI. 
N.O. Cel. (1898) 70 — Type: Koorders 18123 8 (BO), 18124 6 (BO), 18146 8 (BO, L), 18164 6 (BO: 
L, lecto). 


Tree 10-20 m. Twigs in apical portion somewhat flattened but not angular, lower 
down terete, not ridged, 2-5(-10) mm diam., glabrescent from a minute tomentum 
composed of hairs c. 0.1 mm; bark finely striate, brown when older not flaking, 
lenticels smallish, abundant, not very conspicuous. Leaves in two rows, membra- 
nous, oblong-lanceolate to lanceolate, broadest at or slightly above the middle, 
8-23 x 2.5-7.5 cm, base attenuate, top acute-acuminate; upper surface drying 
olivaceous to dark brown, dull, faintly finely paler punctate-pustulate or not (rarely 
with pale, irregularly shaped marks as in H. irya), lower surface early glabrescent, 


70 Gard. Bull. Sing. 38(1) (1985) 


without blackish dots; midrib flat above; nerves 10-15 pairs, flattish and inconspi- 
cuous above, marginal arches not distinct; tertiary venation indistinct, forming a 
rather fine network; petiole 6-16 X 1.5-2.5 mm; leaf bud slender, c. 6-13 x 1-2 mm, 
with hairs c. 0.1 mm long or less. Inflorescences with sparse to dense tomentum of 
hairs 0.1-0.3 mm; in both CO and Q 3-4 times ramified, many flowered, (4-)6-10 x 
4-8 cm, common peduncle 1-2 cm long; bracts pubescent, elliptic to oblong, 2-5 mm 
long, caducous. Flowers in loose clusters of 2-4 each, perianths 2-valved, glabrous 
or in sometimes minutely pubescent at base, pedicels sparsely pubescent with the 
hairs c. 0.1 mm long or less, at base inarticulate. Male perianth + obtriangular to 
transversely ellipsoid, somewhat laterally compressed, 2.2-3.0 x 2.5-4 mm, upper 
part broadly rounded, at base short-attenuate, rather firm, on drying not collaps- 
ing, often bright brown or with a grey-blue tinge; pedicel slender, 1-2 mm long. 
Perianth at anthesis cleft to c. /2-way, valves (0.1-)0.2-0.3 mm thick. Androecium 
transversely ellipsoid or + obtriangular, only slightly or not laterally compressed, 
largely hollow, (1.0-)1.6-2.2 x 1.6-3.0 mm; anthers (18-)20-25, mutually complete- 
ly connate, forming a thin-walled cup, the anthers (sometimes only of one side of 
the androecium) completely inflexed from their middle and reaching nearly to the 
bottom of the cup; free apices of anthers 0-0.1 mm; androphore rather narrow, 
(O-)0.1-0.3 mm long. Female perianth ellipsoid, c. (2.5-)3-3.5 x 2.5 mm, at anthesis 
cleft to c. 4%, valves c. 0.3 mm thick, pedicel 1-2 mm, thinly pubescent with the 
hairs c. 0.1 mm; ovary ovoid-ellipsoid, c. 2-2.3 x 1.5 mm, glabrous, style and 
stigma minute, 2(-3)-lobed, 0.1-0.2 mm long. Fruits 2-10 per infructescence, ellip- 
soid to nearly globose, 1.1-1.6 x 1.0-1.3 cm, glabrous, finely granulate, not or 
hardly tuberculate, drying brown; dry valves 1-1.5 mm thick; stalk 2-4 mm; 
perianth not persisting. 


Distribution. Celebes (incl. Kabaéna Isl.); Moluccas: Ceram, Ambon; running 
wild in the Gardens Jungle of the Botanic Garden, Singapore. 


Cultivated. Java (Bot. Garden Bogor): Forbes 1184a; Rastini 105, (206), (223); Woerjantoro 99. — 
Singapore (Bot. Garden): Ding Hou 134; Murton 149; Ridley s.n., 393; Furtado SFN 34818; Sinclair 
7493. 

CELEBES: b.b. 5432, 8459, 13748, 20754; Elbert 3457; Koorders 18146 fB, 18164 B; Meijer 11286. 

CERAM: Kuswata & Soepadmo 86, 236. 


AMBON: Kuswata & Soepadmo 292; de Vriese & Teysmann s.n.; de Vriese s.n.; Zippel s.n. (“‘mas’’). 


Ecology. Forests; once recorded from sandy loam; 0-600 m alt. Flowers and 
fruits throughout the year. 


Vernacular names. Kolantie, Niniwo (Celebes). 


NOTES 


1. Fieldnotes. Tree without buttresses. Bark smooth or fissured. Wood whitish. 
Flowers yellow, fragrant; anthers yellowish-white. Fruits ramiflorous, yellow to 
light brown. Aril bright red, once recorded as yellow (unripe?). 


2. Variation. The male flowers of Koorders 18146 f (syntype of H. globularia var. 
minahasae), and those of e.g., Ding Hou 134 (Bot. Garden Singapore) are 
relatively large, the perianths being as wide as c. 4 mm. 


New account of Horsfieldia 2 at 


The androecium of b.b. 13748 (N. Celebes) is relatively short, measuring c. 1 Xx 
2.5 mm; its flower are small, c 3 mm in width. 


Kuswata & Soepadmo 236, W. Ceram, has comparatively small fruits, c. 1.1 x 
1.0 cm. 


Elbert 3457 (Kabaéna Isl., limestone; S. Celebes) has rather large fruits, c. 1.6 
cm long, with a distinct pseudo-stalk 1.5-2 mm long. 


3. H. parviflora is easily distinguished by the smooth and rather inflated male 
perianths, which do not collapse on drying, are + obtriangular to transversely 
ellipsoid in lateral view, and usually dry to a bluish or reddish-brown tinge. The 
androecium is largely hollow, inflated, cupshaped, composed of anthers connate 
along the whole length; the distal end from approximately 12-way curved into the 
androecium, almost reaching the bottom. 


4. In Sinclair’s sense, H. parviflora has a much wider circumscription than is 
accepted here. The material he included is here referred to various different species 
such as H. obscurinervia and the variable H. laevigata. 


5. Typification. H. parviflora was described on a female specimen cultivated in 
the Botanic Gardens at Calcutta, and apparently no authentic material is pre- 
served. Sinclair (p. 88, 122) discussed elaborately the identity of Roxburgh’s 
descriptions. 


Although named ‘parviflora’ because of the female flowers, our present species 
has male perianths of 2.5-4 mm wide, which is among the largest in Horsfieldia. 


11. Horsfieldia obscurinervia Merr. Fig. 1A(11) 


H. obscurinervia Merr., Phil. J. Sc. C. Bot. 12, 5 (1917) 265; En. Phil. Fl. Pl. 2 (1923) 182. — Type: de 
Mesa & Magistrado FB 26503 (iso: K). 


Hi. ramosii Merr., Phil. J. Sc. 17, 3 (Sept. 1920) (1921) 254; En. Phil. Fl. Pl. 2 (1923) 182. — Type: 
Ramos BS 35047 (PNH, n.v. (iso: K). 


Tree c. 11 m. Twigs terete, not ridged, 1.5-4 mm diam., glabrescent from a 
minute tomentum of greyish hairs less than 0.1 mm long; bark finely striate, when 
older not flaking; lenticels present, rather distinct. Leaves in two rows, char- 
taceous, oblong-lanceolate to lanceolate, 5-14 x 2-4 cm, broadest at or slightly 
above the middle, base attenuate, tip acute-acuminate; upper surface drying 
olivaceous to brown, + glossy, lower surface early glabrescent, without larger dark 
dots; midrib flat above to slightly raised; nerves 7-15 pairs, slender, very inconspi- 
cuous on both surfaces, marginal arches very inconspicuous; tertiary venation 
hardly visible; petiole 6-15 x 1-1.5 mm; leafbud c. 6-10 x 1.5-2 mm, with hairs 0.1 
mm long or less. Inflorescences sparingly pubescent or subglabrous with hairs c. 
0.1-0.3 mm long; in GC: 2-3 times ramified, rather few-flowered, c. 3-4 x 2-3 cm, 
common peduncle 5-10 mm; bracts and bracteoles not seen, caducous; 9 infloresc- 
ences not seen. Flowers solitary or 2 or 3 together, perianths 2-valved, glabrous; 
pedicel glabrous, at base inarticulate. Male perianth subobtriangular, broadly 
rounded above, + cuneate at base, c. 2 X 2.2 mm, rather firm, on drying not 
collapsing, bright brown, pedicel slender, c. 1 mm long; perianth at anthesis cleft to 
c. ¥2-way, valves 0.2-0.3 mm thick. Androecium + obtriangular or obovoid, 


72 Gard. Bull. Sing. 38(1) (1985) 


narrowed to the base, only slightly laterally compressed, thickish, c. 1.5 X 1.2 mm, 
largely hollow; anthers 11 or. 12 (1.e., 11 or 12 thecae on each side), almost 
completely connate, forming a firm thick-walled cup reaching to c. % of the 
androecium, the anthers at one side deeply inflexed into and almost completely 
filling the cup; free apices of anthers 0-0.1 mm; androphore narrow, short, c. 0.1 
mm long. Female perianths not seen. Fruits 2-5 on once or twice ramified stalk, 
infructescences 2-3 cm long; fruits short-ellipsoid, 1.1-1.3 x 0.9-1.1 cm, almost 
glabrescent but with minute dendroid hairs at base (hence ovary pubescent), finely 
granulate, not tuberculate, drying (reddish) brown; dry valves 1-1.5 mm thick; stalk 
2-5 mm; perianth not persisting. 


Distribution. Philippines: Luzon. 


PHILIPPINES: de Mesa & Magistrado FB 26503; Ramos & Edano BS 33693; Ramos BS 35047. 


Ecology. On low hills at c. 20 m. Flowers in July, fruits November and De- 
cember. 


Vernacular name. Duguan. 
NOTES 
1. Fieldnotes. Small tree; flowers yellow. 


2. Related to H. parviflora from the Moluccas on account of the a!most similar 
flower structure; H. parviflora differs by its larger membranous leaves, larger male 
perianths of 2.5-4 mm width, more (18-25) anthers, thinner-walled and deeper 
androecium-cup, with the anthers usually inflexed at both sides of the androecium, 
and the glabrous ovary and fruit. 


3. Sinclair (p. 83, 90) treated H. obscurinervia as a synonym of H. parviflora. 


12. Horsfieldia ardisiifolia (DC.) Warb. Fig. 1A(12); 9 


Myristica ardisiifolia A. DC., Ann. Sc. Nat. Bot. 4, 4 (1855) 31, t. 4; Prodr. 14, 1 (1856) 203 
(ardisiaefolia). — H. ardisiifolia (DC.) Warb., Mon. Myrist. (1897) 274; Sinclair, Gard. Bull. Sing. 28 
(1975) 3. — Type: Cuming 1702 (iso: L). 


H. warburgiana Elmer, Leafl. Phil. Bot. 3 (1911) 1061; Merr., En. Phil. Fl. Pl. 2 (1923) 183. — Type: 
Elmer 12297 (iso: K & L, only the fruits, see the notes). 


H. gigantifolia Elmer, Leafl. Phil. Bot. 9, 123 (1925) 3120, 3129; 10, 136 (1939) 3763, nom. nud. — 
Type: Elmer 17220 (iso: L). 


Tree 5-10 m. Twigs flattened in the apical part, 2-angular, lower down terete with 
two distinct ridges from petiole to petiole, 3-6(-13) mm diam., early glabrescent 
from the bright rusty tomentum composed of hairs 0.3-0.5(-0.8) mm long; bark 
rather smooth to striate, distinctly lenticellate; not flaking when older. Leaves in 2 
rows, membranous, elliptic-oblong to oblong, 20-40 x 5.5-15 cm, base nearly 
rounded to attenuate, tip acute-acuminate; upper surface drying olivaceous to 
blackish-brown, finely minutely paler pustulate or not, lower surface early glabres- 
cent except for some tomentum remaining on the midrib, consisting of rather 
coarse hairs 0.3-0.5 mm; larger dark dots absent; midrib fairly broad, flattish 
above; nerves 18-28 pairs, slender above, flattish, the marginal arches rather 


Fig. 9. 


Horsfieldia ardisiifolia (A. DC.) Warb. 

a, leafy twig apex, note ridged twig, x 2; b, twig portion with male inflorescence in axil of 
fallen leaf, x %; c, mature male flower, lateral view, X 6; d, ditto, opened, showing 
androecium, X 6; e, androecium, longitudinal section, schematic, X 12; f, mature female 
flower, lateral view, X 6; g, ditto, opened, showing glabrous ovary with minute stigmas, X 6; h, 
twig portion with infructescence with ripe fruits. — a, Ramos BS 39770; b, Sulit PNH 6236; c-e, 
Elmer 12337; f & g, Elmer 17220; h, Conklin PNH 17461. 


73 


74 Gard. Bull. Sing. 38(1) (1985) 


regular and distinct beneath; tertiary venation forming an inconspicuous lax net- 
work; petiole 13-16 x 3-4.5 mm. Leaf bud 10-20 x 3-4 mm, pubescent with hairs 
0.3-0.8 mm long. Inflorescences thinly pubescent with stellate-dendroid hairs c. 0.3 
mm; in ©: 3-4 times ramified, rather many-flowered, broadly pyramidal, c. 7-16 x 
6-14 cm, common peduncle 5-10(-20) mm; in @: 4-8 cm long. Bracts broadly ovate, 
pubescent, c. 3-4 mm long, caducous. Flowers solitary or 2-4 together, perianths 
2-valved, glabrous, pedicels glabrous or glabrescent from sparse hairs, slender, at 
base inarticulate. Male perianth transversely ellipsoid or reniform, moderately 
laterally compressed, drying dull, + collapsed on drying or not, 2.5-3 x 4-4.5 mm, 
above broadly rounded, below broadly rounded to sometimes with a basal sinus; 
pedicel 1-2(-4) mm, glabrous or with few scattered hairs 0.2-0.3 mm; perianth at 
anthesis cleft to c. Y5-%, valves c. 0.2(-0.3) mm thick. Androecium broadly trans- 
versely ellipsoid, slightly laterally flattened, hollow, c. 1.5 x 3-3.5 mm, anthers 
(18-)20-24, for c. '2-way connate and forming a cup in which the anthers from one 
side are deeply inflexed, the anthers from the other side for a large part overarching 
the former; anthers sometimes slightly sagged at base, hiding the rather narrow 
androphore c. 0.2-0.3 mm long. Female perianth subglobose-ovoid, c. 2.5 mm 
diam., at anthesis cleft to c. /2-way, valves c. 0.3 mm thick, pedicel c. 2(-2.5) mm 
long, (sub)glabrous; ovary broadly ovoid-subglobose, 1.5-1.7 mm diam., glabrous, 
stigmas sessile, as 2 minute lobes c. 0.1-0.2 mm. Fruits 2-6 per infructescence, 
broadly ellipsoid to subglobose, 20-25 x 17-20 mm, glabrous (or possibly with few 
minute hairs at base), finely rugulose, without marked tubercles, drying (reddish- 
)brown; dry valves 1.5-2 mm thick; stalk 3-6 mm long; perianth not persisting. 


Distribution. Philippines: Luzon, Mindoro, Sibuyan, Samar, Leyte. 


PHILIPPINES: Brass 1220; BS (Bermejos) 1518, (Ramos) 39770, 40823, 46414; Cuming 1702; Elmer 
7094, 12067, 12297, 12337, 17220; For. Bur. (Parras & Aduviso) 28297; Gaudichaud s.n.; PNH 
(Celestino & Castro) 1925, (Sulit) 6236; (Conklin) 17461. 


Ecology. Lowland forest in moist valleys. Flowers and fruits throughout the year; 
0- c. 400 m. 


Vernacular names. Aragay (Mangalang, Mindoro), Dagoan (C. Biscuay), Tapol 
(Tagbilaran), Lagasi (Biscuay Isl.). 


NOTES 


1. Fieldnotes. Flowers reported as yellow or lemon yellow, fragrant. Fruits 
orange-red. 


2. According to Sinclair (p. 5) this species is closely related to H. spicata. 
Sinclair’s idea of H. spicata embodied several taxa, which are presently regarded as 
different species because of differences in the androecium. As can be seen from the 
present key to the species it is now considered as being close to species like H. 
parviflora and H. smithii from the Moluccas, i.e., of the group with anthers strongly 
incurved or inflexed into the androecium-cup. H. ardisiifolia is quite distinct by its 
stout habit, with thick twigs which are winged or ridged, large leaves, coarse 
tomentum on the leaf bud, large male perianths (4-4.5 mm wide), broad 
androecium with the anthers deeply incurved and clasping each other. 


3. Of the type of H. warburgiana, Elmer 12297, | have seen only two isotypes, in 
K and L. They are conspecific and consist of a leafy twig, and fruits in an attached 


New account of Horsfieldia 2 75 


envelope. The fruits belong to H. ardisiifolia, but the leafy twigs are most likely H. 
macrocoma, a species which is recently referred by me to a new genus Endocomia 
(1984). 


In 1959 Sinclair identified Elmer 12297 in L as H. ardisiifolia but later he 
regarded that name a synonym of H. brachiata var. sumatrana, a taxon of which the 
Philippine specimens are presently referred by me to various other species. 


13. Horsfieldia talaudensis de Wilde, sp. nov. Fig. 1A(13) 


Horsfieldia species perianthiis masculis 2-valvatis atque antheris inflexis, ex affinitate H. ardisiifoliae, 
ab ea differt virgis teretibus non-angularibus, perianthio minore c. 2.5 mm diam. atque pedicellis 
pubescentibus. Typus: Lam 2628 (L). 


Tree 15-35 m. Twigs terete, not ridged, towards the apex 2-4(-7) mm diam., early 
glabrescent, tomentum scarce, composed of hairs 0.1 mm or less; bark striate, not 
flaking when older, lenticels rather inconspicuous. Leaves in 2 rows, membranous 
to chartaceous, oblong-lanceolate, 8-30 x 2.5-10 cm, base attenuate, tip acute- 
acuminate, finely paler pustulate on both surfaces; upper surface drying greenish to 
dark brown, lower surface bright brown, early-glabrescent, hairs minute, stellate- 
scaly, 0.1 mm long or less, dark dots absent; midrib flat above; nerves 12-20 pairs, 
above slender, flat, inconspicuous, marginal arches not distinct, tertiary veins thin, 
indistinct on both surfaces; petioles 10-18 x 1.5-3 mm, leaf bud c. 10 X 2 mm, with 
hairs c. 0.1 mm. Inflorescences densely pubescent with hairs 0.1-0.2 mm, c. 3 times 
ramified, in & and Q c. 4-8 x 3-4 cm, rather many-flowered, common peduncle 
1.5-2 cm; bracts broadly ovoid-ellipsoid, densely pubescent, 1.5-3 mm long, cadu- 
cous. Flowers in © in clusters of 2-3 each, perianths 2-valved, glabrous; pedicel 
pubescent, at base inarticulated. Male perianth transversely ellipsoid, only little 
laterally compressed, c. 2-2.2 X 2.5-3 mm, broadly rounded above and at the base, 
glabrous, drying brown, not collapsing; pedicel slender, 1-1.5 mm, rather densely 
pubescent with grey or pale brown hairs c. 0.1 mm; perianth at anthesis cleft to 
*/3-Y5, valves rather firm, c. 0.2-0.3 mm thick. Androecium transversely ellipsoid to 
reniform, not much compressed, c. 1.5 X 2 mm; anthers c. 18, closely set, connate 
for about ’2-way and forming a + saucer-shaped cup or cavity into which the free 
apical halves of the anthers are inflexed; young anthers septate; androphore 
slender, 0.2-0.3 mm long. Female flowers not seen. Fruits 3-10 per infructescence, 
short-ellipsoid, top and base rounded, 1.5-1.6 x 1.3-1.4cm, glabrescent, tomentum 
of stellate hairs c. 0.1 mm or less (hence ovary pubescent), drying brown, without 
conspicuous tubercles, dry valves c. 1.5 mm thick, stalk 3-4 mm long; perianth not 
persisting. 


Distribution. Moluccas: Talaud Isls. (Karakelong); possibly Celebes: Minahassa 
(see notes). ; 
CELEBES. Minahassa: Koorders 18136 8 (doubtful, see notes). 


MOLUCCAS. Talaud Isls.: Lam 2638, 2650, 2811, 2929. 


Ecology. Old forest on mountain slopes; 70-200 m alt. Flowers in April, fruits in 
April and May. 


Vernacular name. Laran’a. 


76 Gard. Bull. Sing. 38(1) (1985) 


NOTES 
1. Fieldnotes. Tree to 35 m. Ripe fruits orange or brownish yellow. 


2. Possibly endemic on the Talaud Isls. A species of the group with the 
androecium having strongly inflexed anthers, characterised by the firm, transverse- 
ly ellipsoid to subglobose male perianth, by the anthers connate to c. 12-way, by 
hairy, short but slender pedicels, and by the pubescent ovary (thinly pubescent 
young fruit). 


3. The specimen from the Minahassa (Celebes; Koorders 18136 8) is sterile but 
agrees vegetatively, in the leaf colour and texture, and very well with the fruiting 
specimens from Talaud Isls. 


4. The specimens belonging to the present new species were included by Sinclair 
in H. parviflora, a species presently accepted in a much narrower sense. 


14. Horsfieldia samarensis de Wilde, sp. nov. Fig. 1A(14) 


Horsfieldia species perianthiis masculis 2-valvibus et antheris profunde inflexis, ex affinitate H. 
ardistifoliae atque H. talaudensis, ab H. ardisiifolia differt virigis non-angularibus et floribus minoribus, 
ab H. talaudensi antheris inflexis solum ad umum androecii latus atque pedicellis perianthio longioribus. 
— Typus: Gutierrez PNH 147374 (L). 


Tree 5 m. Twigs terete, not ridged, towards the apex 1.5-3.5 mm diam., early 
glabrescent, tomentum minute, of greyish hairs less than 0.1 mm; bark finely 
striate, when older not flaking, lenticels rather inconspicuous. Leaves in 2 rows, 
membranous, oblong-lanceolate to lanceolate, c. 7-11 x 2-3 cm, base attenuate, 
tip acute-acuminate, not finely, paler pustulate: upper surface drying dull olli- 
vaceous, lower surface bright brown, early glabrescent, hairs sparse, less than 
0.1 mm, without dark dots; midrib slightly raised above; nerves 10-13 pairs, above 
slender, flat, rather contrasting in colour; marginal arches on lower surface faint but 
rather regularly looping; tertiary venation inconspicuous on both surfaces; petioles 
c. 8-10 x 1-1.5 mm, leaf bud c. 10 X 1 mm, with hairs less than 0.1 mm. 
Inflorescences glabrescent or thinly pubescent by stellate scales c. 0.1 mm or less, 
rather slender, in CG’: 2-3 times ramified, 3-4 xX 1.5-2.5 cm, rather few-flowered, 
common peduncle c. 1-1.5 cm; bracts not seen, caducous. Male flowers solitary or 
in loose clusters of 2 or 3 together; perianths 2-valved, glabrous, pedicels glabrous, 
at base inarticulate. Male perianth transversely ellipsoid, only slightly laterally 
compressed, 2-2.2  2.5-2.7 mm, above and below broadly rounded, drying brown, 
firm, not collapsing, glabrous; pedicel slender, (1.5-)2-3 mm long, glabrous; 
perianth at anthesis cleft to 73-%, the valves rather firm, 0.2(-0.3) mm thick. 
Androecium broadly obovoid to transversely short-ellipsoid, 1.2-1.3 x 1.4-1.5 mm, 
thickish, 0.8-0.9 mm thick; anthers 14 or 15, closely set, septate when immature, 
largely connate and forming a rather thick-walled saucer-shaped cup into which the 
anthers at one side inflect deeply nearly to the base, clasping and covering the 
other anthers, the inflexed parts of the anthers mutually free; androphore narrow, 
(0-)0.1 mm long. Female flowers and fruits not seen. 


Distribution. Philippines: Samar Isl. (only known from the type). 


Ecology. North slope, 600-800 ft. Flowers in May. 


New account of Horsfieldia 2 77 


NOTES 
1. Fieldnotes. Tree c. 5 m tall, dbh c. 6 cm. Inflorescence (flowers) green. 


2. According to the flower structure related to H. talaudensis, but differing in 
several points. H. talaudensis is stouter, with male inflorescences stouter, more 
densely pubescent; its pedicels are shorter (shorter than the perianth) and densely 
pubescent, the perianth somewhat larger, the androecium with the anthers inflexed 
from both sides into the cavity. 


3. The specimen on which the present new species is based was collected after 
Sinclair’s revision. 


15. Horsfieldia smithii Warb. Fig. 1A(15) 


H. smithii Warb., Mon. Myrist. (1897) 270, t. 21, 1-3. — Myristica smithii (Warb.) Boerl., Handl. FI. 
Ned. Ind. 3, 1 (1900) 87 — Type: Smith s.n. (1797, in BM sheet numbered 296) (iso: BM, K, L; 
LINN, B+, BR & G, n.v.). 


Tree 10-20 m. Twigs in apical portion flattish and 2-angled, often somewhat 
yellowish, lower down subterete, with two distinct or faint ridges or lines from 
petiole to petiole, 2.5-5(-8) mm diam., glabrescent, minute tomentum with hairs 
0.1 mm long or less; bark finely striate, when older not flaking, lenticels rather 
small and inconspicuous. Leaves in 2 rows, membranous, oblong-lanceolate, (10-) 
15-30 Xx (4-) 5-10 cm, base attenuate, tip acute-acuminate; upper surface drying 
olivaceous to brown, with fine minute paler pustules or not, almost always with 
larger irregular whitish marks, lower surface early glabrescent, hairs 0.1 mm long 
or less; without dark dots; midrib above slender, flat; nerves 10-18 pairs, above thin 
and flat or slightly raised, inconspicuous, marginal arches not distinct; tertiary 
venation forming a rather lax network, indistinct; petiole 10-16 x 1.5-2.5 mm; leaf 
bud c. 10 X 2 mm with hairs c. 0.1 mm or less. Inflorescences with sparse to dense 
tomentum of hairs c. 0.1 mm or less; in CG: (2-) 3-4 times ramified, many- 
flowered, c. 5-8 X 4-8 cm; in 9: c. 2-3 cm long; common peduncle 5-15 mm; bracts 
not seen, caducous. Flowers 2-4 together, perianths and pedicels glabrous, 
perianths 2-valved; pedicel glabrous, at base inarticulate. Male perianth subcircular 
to distinctly transversely ellipsoid or slightly reniform, laterally compressed, dull 
and usually + collapsed on drying, 2.5-3.0 X 3-4 mm, above broadly rounded, at 
base rounded, or subtruncate, or shortly tapering; pedicel slender, 1.5-2 mm; 
perianth at anthesis cleft to c. 74-%4, valves 0.1-0.2 mm thick. Androecium as 
viewed laterally from the broad side transversely ellipsoid, inflated, consisting of a 
bunch of anthers 1-1.5(-2.0) x 2.5-3.5 mm, and a sterile basal part or androphore, 
rather tapering, 0.5-0.8 mm long; anthers 12-15 (i.e., 12-15 thecae at both sides of 
the androecium), the thecae slender, mutually almost free, c. 1.5-2 mm long, their 
upper halves deeply curved into the cavity, which extends almost to the base of the 
androecium, the thecae subdorsally attached to the rim of the androphore only at 
the base. Female perianth ovoid-ellipsoid, 2-3 mm diam., at anthesis cleft to c. 
Y2-way, valves c. 0.3 mm thick, pedicel c. 2 mm long, thinly pubescent; ovary 
globose-ovoid, c. 2 X 1.7 mm, glabrous, stigma c. 0.1 mm long or less. Fruits 1-3 
per infructescence, ellipsoid, 1.5-2.0 (-3.0) x 1.4-1.6 cm, glabrous, drying brown, 
with scattered small tubercles or lenticels; dry valves 1.5-2 mm thick; stalk 3-5 mm 
long; perianth not persisting. 


78 Gard. Bull. Sing. 38(1) (1985) 


Distribution. Moluccas: Ceram, Banda, Damar Isls., possibly Ternate (see 
notes). 


CERAM: Kornassi (exp. Rutten) 2/8; Rutten 1776. 


BANDA: Cult. hort. Bogor: Anon. 270 (anno 1901); Forbes 1158 (anno 1880); Koerniasih 42; Banda, 
Chr. Smith s.n. & 296 (May 1797). 


DAMAR Isls.: Riedel s.n. (syntype H. novoguineensis, not the lectotype). 


TERNATE: ? Smith s.n. (see the notes). 
Ecology. Nothing known. 


Vernacular names. Pohon lobi-lobi (Banda, Ceram), Pala oetan (Ceram). 


NOTES 


1. Fieldnotes. Flowers yellow, in May and October; fruits yellow with whitish 
dots, in October; 0-100 m alt. 


2. Closely related to H. palauensis, H. parviflora and H. ardisiifolia, with which 
it shares the character of largely free thecae which are curved into the hollowed 
+ cup- or saucer-shaped androphore; the more or less incurved anthers of H. irya 
are reminiscent, but in this latter species the perianth is much smaller and the 
anthers are shorter and free at the apex for only c. 0.2 mm; in H. moluccana the 
male perianth is + pear-shaped and the anthers are largely inter-connate. 


3. The present species as well as related species like H. parviflora, H. ardistifolia, 
and also H. irya and H. moluccana may have somewhat ridged or angular twigs, 
and in sterile or fruiting stages may be confused with species with typically ridged or 
winged twigs, e.g., H. angularis, and a few species confined to West Malesia. 


4. With Warburg’s key, H. smithii is keyed out (p. 262) in a group (series Smithii) 
which is characterized by 2-valved perianths, ridged twigs and interconnate 
anthers. However, in reality the anthers appear to be almost entirely or largely 
mutually free, with the long free portions of the thecae deeply curved into the + 
bowl-shaped androecium. 


5. Sinclair included the present species in his broad concept of H. spicata, the 
latter presently accepted by me in a much narrower sense, and regarded as distinct 
by the spicate male and female inflorescences, pear-shaped male perianths, erect 
uncurved anthers, etc. 


6. The specimen from Damar Isls. (Riedel. s.n.) was determined by Sinclair as H. 
parviflora which in his notion is also a broad species and which I accept in a 
narrower sense. H. parviflora differs from H. smithii by the less distinctly ridged or 
lined twigs, the thicker male perianths which do not collapse on drying, the 
more rigid and broader (wider) androecium with the thecae only mutually free in 
the long incurved portions, and by the absence of irregularly shaped whitish marks 
on the leaves. 


7. The specimens from Ternate (possible collected by Smith, Oct. 1801; in BM). 
bears immature female flowers with glabrous ovaries. The indumentum of the 
inflorescences is rather thick and woolly, and the specimen could well be H. irya, a 
species usually with similar whitish markings on the leaves. 


New account of Horsfieldia 2 79 


16. Horsfieldia palauensis Kanehira Fig. 1A(16) 


H. palauensis Kanehira, New Trees Micronesia I, in Bot. Mag. Tokyo 46 (1932) 452 (‘palauense’); FI. 
Micronesica (1933) 111, Fig. 33; Bot. Mag. Tokyo 47 (1933) 670, in the notes to H. amklaal; An 
Enumeration of Micronesian Plants, in J. Dept. Agric. Kyushu Imp. Univ. 4, 6 (1935) 319. — Type: 
Kanehira 270 (Q), Palau Isls. (FU, n.v.; iso: BISH, FI; NTS, n.v.). 


H. glabra auct. non (Bl.) Warb.: Kanehira, Bot. Mag. Tokyo 45 (1931) 280. 


Tree 7-15 m. Twigs terete, lined from petiole to petiole, c. 3-4(-8) mm diam., 
early glabrescent, tomentum minute, with hairs c. 0.1 mm; bark striate, when older 
not flaking, lenticels minute and rather indistinct. Leaves in 2 rows, membranous to 
subchartaceous, oblong(-lanceolate) to lanceolate, 10-22 x 3.5-7 cm, base short- 
attenuate to rounded, tip obtuse to acute-acuminate; upper surface drying oli- 
vaceous brown to dark brown, not finely pustulate, sometimes with a few paler 
marks of irregular shape, lower surface early glabrescent from hairs 0.1 mm or less, 
without dark dots; midrib slender above, flat or slightly raised; nerves 10-20 pairs, 
thin and flat above, inconspicuous, marginal arches indistinct; tertiary venation 
forming a rather lax network, indistinct or invisible; petiole 10-16 x 1.5-2 mm; leaf 
bud c. 10 X 2 mm, with tomentum of hairs c. 0.1 mm or less. Inflorescences with 
sparse to dense woolly rusty tomentum with hairs 0.1-0.3 mm; in O and Q: 2-3 
times ramified, rather few- to many-flowered, + short, c. 3-6 X 2-4 cm; common 
peduncle 5-15 mm; bracts broadly ellipsoid, 2-3 mm long, caducous. Flowers 3-6 
together, perianths glabrous, 2-valved, pedicels glabrescent or with sparse to + 
dense tomentum of hairs 0.1-0.2 mm, inarticulate at base. Male perianth sub- 
obtriangular to transversely ellipsoid or kidney-shaped, rather distinctly laterally 
compressed, dull dark brown, c. 1.7-2.0(-2.3) xX 2-2.5(-3.0) mm, above broadly 
rounded, at base subtruncate to short-cuneate; pedicel 0.5-1.0(-1.5) mm; perianth 
at anthesis cleft to c. 73-%, valves c. 0.2-0.3 mm thick. Androecium subellipsoid, c. 
1.0-1.5 x 1.8-2.0(-2.5) mm, with a sterile basal part mainly consisting of the 
androphore, + hollowed out and rather tapering, c. 0.5-0.6 mm long; anthers c. 
8-13, i.e., c. 16-26 thecae and these mutually free for the upper *4 or more, c. 
0.7-1.5 mm long, strongly incurved, the ones at one side of the androecium usually 
covering those of the other side. Female perianth broadly ellipsoid, c. 2.5-3.0(-3.5) 
x 2.2-3.0 mm, at anthesis cleft to c. '2-way, valves c. 0.3 mm thick, pedicel 1-2 mm 
long, + sparsely pubescent with hairs c. 0.1 mm; ovary globose-ellipsoid or obo- 
void, c. 1.6-2.0 mm long, glabrous, stigma very minutely 2-lobed. Fruits 2-5 per 
infructescence, ellipsoid, 2.5-3.0 x 1.5-2.0 cm, glabrous, drying dark brown, with- 
out tubercles; dry valves c. 2-3 mm thick; stalk 2-5 mm; perianth not persisting. 


Distribution. Caroline Isls.: Palau Isls. 


PALAU ISLS.: Hosokawa Takahide 6756; Kanehira 270, 1847, 1958, 1960, 2371; Masahiko Takamat- 
su 1205, 1668, Shearard & Spence 94; Tuyama s.n., 10 Sept. 1937, 9, 14, 15, 17, 19, s.n. Aug. 1939. 


Ecology. Locally abundant in primary lowland forest, usually not in the wetter 
localities and usually at somewhat higher altitudes; apparently not or only rarely in 
mangrove forest. Flowers and fruits throughout the year. 

Uses. Pericarps edible, eaten by the natives. 


NOTES 


1. Fieldnotes. Flowers recorded as yellow or orange. 


80 Gard. Bull. Sing. 38(1) (1985) 


2. Closely related to H. smithii from the Moluccas, which differs chiefly by the 
larger male perianth, c. 3-4 mm wide, by the anthers which are all (i.e., not only 
from one side of the androecium) incurved towards the centre of the androecium, 
and by the generally somewhat thinner leaves with the lateral nerves on the upper 
surface flat or slightly raised. In H. palauensis the older leaves are somewhat 
chartaceous, above with the lateral nerves flat or sunken, and the tertiary venation 
very indistinct or even invisible. 


3. Sinclair (pp. 112, 119, 124) treated the present species as a synonym of H. 
spicata. However, as understood by Sinclair, H. spicata appears to be a heter- 
oOgeneous species which is in my present treatment divided among various different 
species. H. spicata, in the restricted and original sense, is a species presently 
regarded as confined to Celebes and the Moluccas. 


4. As far as I know, only two species occur on the Palau Isls., viz. the wide- 
spread H. irya (syn. H. amklaal, H. nunu), and H. palauensis, an endemic species 
closely related to H. smithii from the Moluccas. 


5. Doubtful specimens. Two sheets of Takamatsu 1205, in BISH, one female 
flowering and the other a fruiting specimen, somewhat deviate by the membranous 
leaves with rather distinct, slightly raised lateral nerves and venation. 


17. Horsfieldia olens de Wilde, sp. nov. Fig. 1A(17) 


Horsfieldia species perianthio masculo glabro fere ad basin 3-valvi, androecio elongato, ex affinitate 
H. sepikensis, ab ea differt virgis angularibus, perianthio minus elongatis, atque antheris haud profunde 
inflexis. — Typus: NGF 31966 (L). 


Tree 10-35 m. Twigs in apical portion distinctly quadrangular through ridges 
from both sides of the bases of the petioles, lower down stem merely ridged or 
lined, 2-4(-8) mm diam., early glabrescent from greyish to brown tomentum of 
hairs c. 0.1 mm; bark striate, when older not flaking, lenticels usually distinct. 
Leaves in 2 rows, chartaceous, oblong (to oblong-lanceolate), broadest at or above 
the middle, 7-14 x 2.5-6 cm, base attenuate, tip either rounded, or obtuse, or 
bluntly short-acute-acuminate, upper surface drying brown to blackish, without or 
with few whitish minute dots or pustules, lower surface early glabrescent, hairs 
minute, c. 0.1 mm or less, without blackish dots but irregularly shaped pustules of a 
different nature present; midrib raised above; nerves 7-10 pairs, above and 
below thin, flattish, and inconspicuous, the submarginal arches rather regularly 
shaped; tertiary venation forming a lax network, very faint on both surfaces; 
petioles relatively long and slender, 12-20 * 1.5-2 mm; leaf bud 6-10 X c. 1.5 mm, 
with hairs c. 0.1 mm long. Inflorescences glabrescent or with sparse tomentum of 
stellate hairs c. 0.1-0.2 mm, rather short and stout, 2-3 times ramified, common 
peduncle 1-10 mm long, rather many-flowered, in C&’ and @ (according to the 
infructescences): 2-6 xX 1.5-4 cm, bracts not seen, caducous. Flowers in loose 
clusters of 3-5(-7) together; perianths 3-(or 4-) valved, glabrous; pedicels slender, 
glabrous, at base + articulated or not. Male perianth broadly ellipsoid to globose, 
not angular, c. 1.8-2.3 mm diam., top and base rounded, pedicel not tapering, 2.5-4 
mm long, glabrous; perianth at anthesis cleft to c. % or nearly to the base, valves c. 
0.2 mm thick. Androecium + obovoid, blunt-triangular, c. 1.5 x 1.0 mm (hence 
not completely filling the perianth); anthers c. 10-12, free apices c. 0.3-0.8 mm, 
incurved, those of one side clasping the others; column rather broad and solid, 
hollowed for the upper 4 to 4; androphore rather broad, up to 0.1 mm long. 


New account of Horsfieldia 2 81 


Female flowers not seen. Fruits 2-6 per infructescence, ellipsoid, 1.0-1.6 x 0.8-1.2 
cm, top minutely pointed, base sub-attenuate, glabrous, without or with sparse 
small tubercles or lenticels, dry valves c. 1-1.5 mm thick; stalk 1-4 mm long; 
perianth not persistent. 


Distribution. New Guinea: Irian Jaya, Digul (SE. New Guinea); Papua New 
Guinea, Western Prov. 


NEW GUINEA. Irian Jaya: (Bouman) BW 3234, (Nautje) 6530, 6608; Soegeng 413. — Papua New 
Guinea, Western Prov.: Lae 51821; NGF 8297, 31770, 31966. 


Ecology. Swamp edges, in fringes (with Acacia) of savanna and rain-forest; ridge 
forest, primary forest on level land inundated in the wet season, swampy forest on 
peaty soil; also in Myrtaceae-Vatica-Campnosperma forest on well-drained podsol- 
ground; 0-200 m alt. Flowers in June, fruits from February to August. 


Vernacular names. Isasir, Jisasir, Jesaser (Asmat lang.), Selamae (Kunga dial.), 
Ma-tak (Kinuga Dist.). 


NOTES 


1. Fieldnotes. Once reported to have small stilt roots. Bark longitudinally fis- 
sured, brown or red-brown, or blackish brown, inside reddish, with some reddish 
exudate; reported to have a very offensive smell, or a strong disinfectant smell. 
Wood whitish to yellow. Leaves + leathery, once recorded as bluish-green above. 
Flowers yellow. Fruits orange-yellow or orange, seeds orange or red. 


2. Apparently closely related to H. sepikensis, one of the few other New Guinean 
species with 3-merous perianths. That species differs in its non-angular twig-apices, 
more elongate perianth, more slender inflorescences, membranous leaves, and 
apparently a different ecology. The present new species is mostly found in dry or 
wet habitats on poor peaty or podsolic soils. 


3. Specimens of the present species were included by Sinclair (p. 125, 126) in H. 
spicata var. sepikensis (Mkgf.) Sinclair, a taxon which appears to be a heterogenous 
entity. 


18. Horsfieldia sepikensis Markgraf Fig. 1A(18); 10 


Horsfieldia sepikensis Markgraf, Bot. Jahrb. 67, 2 (1935) 147. — Horsfieldia spicata var. sepikensis 
(Mkgf) Sinclair, Gard. Bull. Sing. 28 (1975) 125, p.p. — Type: Ledermann 8016 (B*; iso: SING, 
n.v.). 


Tree 10-25 m. Twigs terete, faintly ridged or not, towards apex 2-4(-6) mm 
diam., early glabrescent, tomentum of hairs greyish brown, c. 0.1 mm long or less; 
bark striate, when older not flaking, lenticels rather small but distinct. Leaves in 2 
rows, membranous, oblong to oblong-lanceolate, broadest usually at or above the 
middle, 8-17 x 3.5-6 cm, base attenuate, tip acute-acuminate with the very tip 
usually bluntish; upper surface drying dark brown, without minute whitish pus- 
tules, lower surface early glabrescent, minute hairs less than 0.1 mm, without 
blackish-brown dots; midrib above flattish or slightly raised; nerves 8-12 pairs, 
above thin and flat; tertiary veins forming a lax network, very faint on both 
surfaces; petiole relatively long, 15-24 x 1-2 mm, leaf bud c. 10 k 1 mm, with hairs 


82 


ere 


Horsfieldia sepikensis Markgraf 

a, twig portion with male inflorescence, X '2; b, mature male flower, lateral view, Xx 12; c, 
ditto, opened, showing androecium, Xx 12; d, androecium, longitudinal section, schematic, x 
12; e & f, portions of female flowering twig, inflorescences axillary to leaves and fallen leaves, 
x 12; g, mature female flower bud, x 12; h, ditto, at full anthesis, showing glabrous ovary and 
large broadly 2-lipped stigmas, x 12. — a-d, from Hoogland & Craven 10255; e-h, Hoogland 
& Craven 10237. 


New account of Horsfieldia 2 83 


less than 0.1 mm. Inflorescences glabrescent or with sparse hairs c. 0.1 mm, 2-3 
times ramified, many-flowered, in CG: 7-12 x 4-6 cm, in Q: 2-4 x 1.5-2 cm, 
common peduncle c. 0.5-1.5 cm long; bracts not seen, caducous. Flowers in loose 
clusters of 2-7 together; perianths 3-(or 4-) valved, finely punctate, in mature bud 
rather angular (though perianths never tightly clustered), glabrous; pedicels glab- 
rous, at base not articulated. Male perianth broadly ellipsoid-obovoid, + 3- 
angular, c. 1.5-2.0 x 1.4-1.6 mm, at apex acutish, base + attenuate, pedicel 
slender, not tapering, 2-3 mm long; perianth at anthesis cleft to c. %, valves c. 0.1 
mm thick. Androecium + obovoid, + bluntish 3-angular, c. 1.5 x 1-1.2 mm; 
anthers 12-14, tightly set, septate before maturity, free apices c. 0.2-0.5 mm long 
and these + curved into the hollowed upper '3 part of the rather broad anther 
column (anthers at one side of the androecium in Hoogland & Craven 10255 
mutually touching each other in a fish-bone pattern, see fig. 10) androphore 0-0.1 
mm, rather narrow. Female perianth ellipsoid, c. 2.0 x 1.6-1.8 mm, 3- or 4-valved, 
cleft at anthesis almost to the base, valves c. 0.1-0.2 mm thick, pedicel 1.5-2 mm 
long; ovary ovoid, c. 1.5-1.8 X 1.2-1.5 mm, glabrous, style absent, stigmas relative- 
ly very large, consisting of two broad fleshy lobes c. 1.0 X 0.2 mm, only c. 0.1 mm 
high. Fruits not seen. 


Distribution. Papua New Guinea: East Sepik Prov. 


PAPUA NEW GUINEA. East Sepik Prov.: Hoogland & Craven 10237, 10255; Ledermann 6738. 


Ecology. Primary and secondary forest, riverine forest; ridge forest; 0-50 mm. 
Flowers and fruits throughout the year. 


Vernacular name. Bangera (Waskuk lang., Sepik). 
NOTES 
1. Fieldnotes. Flowers yellow. 


2. A noteworthy species because of its predominantly 3-merous flowers, the only 
other species with 3- or 4-merous flowers in New Guinea being the + related H. 
olens. Furthermore, the present species stands apart by its thick, conspicuous 
stigmas. It is in many cases reminiscent of a species of Gymnacranthera, but the 
New Guinean G. paniculata var. zippeliana differs in the nature of the hairs (on leaf 
bud, and flowers), by the hairy perianths, split at anthesis to only c. /2-way deep, 
by a hairy ovary, and a different texture and colour of the leaves (whitish below). 


3. The fruits are reported by Markgraf in the original description as globose, c. 
13-15 cm diam., the dry pericarp c. 2 mm thick; seed globose, c. 1 cm diam. I have 
not seen fruits of our present species, and the fruiting Ledermann-specimens 
formerly in the Berlin herbarium have probably all been destroyed. 


4. | have not seen the isotype Ledermann 8016 (C), in SING. The flowers are 
described by Markgraf in the original description as being cleft to c. /2-way deep at 
anthesis. However, in the male and female specimens I saw have the perianths 
split to almost the base. Ledermann 6738, in K, a duplicate of one of the authentic 
collections cited by Markgraf, has good male flowers. See further under note 5. 


5. Sinclair, who examined a duplicate of the holotype Ledermann 80/6 in SING, 


84 Gard. Bull. Sing. 38(1) (1985) 


accepts the present species as a variety under H. spicata, including specimens with 
2-valved as well as with 3-valved perianths. In my opinion the 3-merous perianths 
are typical for the present species, which are endemic in the Sepik area, one which 
has no close relationship with H. spicata from the Moluccas. 


6. Hoogland & Craven 10255 contains one flower with a ‘double’ androecium; 
the perianth is somewhat larger and has 4 valves. Female flowers are either 3- or 
4-valved. 


19. Horsfieldia sylvestris (Houtt.) Warb. Fig. 1A(19); 11 


Myristica sylvestris Houtt., Nat. Hist. 2, 3 (1774) 340 — Horsfieldia sylvestris (Houtt.) Warb., Mon. 
Myrist. (1897) 337, t. 22 fig. 1-6; Sinclair Gard. Bull. Sing. 28 (1975) 142. — Type: not indicated. 


M. salicifolia Willd. in Roem. & Usteri, Mag. Bot. 3, 9 (1790) 26; Sp. Pl. (4th ed.) 4, 2 (1806) 871; 
Roxb., Fl. Ind. ed. Carey (1832) 846 — Type: not known. 


M. pinnaeformis Zipp. (msc.) ex Miq., Ann. Mus. Bot. Lugd.-Bat. 2, 1 (1865) 49 — Type: Zippelius s.n. 
(180) (L). 


M. pendulina Hook.f., Fl. Brit. Ind. 5 (1890) 859; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 329, pl. 
170 — Type: Cantley s.n. (A, CAL, n.v.; K). 


M. edulis F.v.M., in sched. (Hb. v. Miller, d’Alberis 11, MEL, not seen). 


H. sylvestris var. villosa Warb., Mon. Myrist. (1897) 341 — Type: Beccari 696 (Fl, n.v.), Warburg 20708 
CA;, Bt, 'n.v2): 


Tree 7-40(-60) m. Twigs stout, terete, hollow, in young innovations when dry + 
angular or flattish, usually thinly ridged, 4-14(-20) mm diam., glabrescent from 
rusty + woolly tomentum composed of hairs 0.3-1.0(-1.5) mm; bark faintly striate, 
when older not flaking, with coarse lenticels. Leaves in 2 rows, (thinly) char- 
taceous, lanceolate to lanceolate-linear, parallel-sided, (17-)20-45 x 3-7(-9) cm, 
base rounded to short-attenuate, tip long acute-acuminate; upper surface drying 
usually dull, greenish brown to dark brown, minutely pustulate or not, lower 
surface late glabrescent or with (partially) persistent or sub-persistent tomentum of 
mixed hairs 0.1-1(-1.5) mm; without dark brown dots; midrib flattish above, 
late-glabrescent; nerves 30-42 pairs, thin, above flat or sunken, beneath with 
distinct marginal arches; tertiary venation forming a lax network, distinct (and then 
the leaves + bullate) or usually not distinct above; petiole short, 2-7 x (2-)3-5 mm, 
usually shortly winged, the lamina being decurrent; leaf bud generally stout, up to 
8 cm long, densely woolly-pubescent. Inflorescences pubescent or late-glabrescent, 
hairs woolly, 0.5-1 mm long, in CO: large, many-flowered, 3-5 times ramified, 
paniculate, 7-20(-30) x 4-10(-14) cm, in 9: 4-10(-15) cm long; common peduncle 
2-7 cm X 2-5 mm, at base with a few persistent bluntish cataphylls 2-4 mm long; 
bracts rather late-caducous, + concave, (2-) 4-8(-16) mm long. Flowers in C in 
loose clusters of 4-10 each, in 2 up to 5 only; perianths 2-(or 3-) valved, often 
somewhat angular, glabrous or at base glabrescent, pedicels slender, glabrescent or 
with persistent tomentum of hairs c. 0.3 mm, at base inarticulate; flowers before 
anthesis, especially in 0’, densely packed into subglobose or ellipsoid glomerules 
4-7 mm diam. wrapped in bracts. Male perianths obovoid to narrow-obovoid, or 
clavate, irregularly shaped and angular by being closely packed in bud, c. 1.5-2.1 x 
Q.5-1.3(-1.5) mm, at apex obliquely obtusish, towards base usually + tapering into 
pedicel c. 0.2-1.5(-2)  c. 0.3 mm; perianth at anthesis split to c. %3-'2-way, valves 
0.1-0.3 mm thick, sometimes with a few pale dots. Androecium ellipsoid-oblong, c. 


Mme he, 
fae 


~ 
4 
x 
s 
Vy 
\ 


Fig. 11. 


a bss 


Horsfieldia sylvestris (Houtt.) Warb. 

a, leafy twig apex, x 2; b & c, twig portions with respectively immature and full-grown male 
inflorescences, note bracts in b, X %; d, male flower, X 6; e, ditto, opened, showing 
androecium, X 12; f, twig portion with female inflorescence axillary to fallen leaf, x 2; g, 
female flower, lateral view, opened, showing glabrous ovary, note much larger size as 
compared with the male flowers, x 6; h, twig portion with infructescence with mature fruits. 
— a, f, g, de Vogel 3069; b, de Vogel 3094; c-e, Craven 739; h, de Vogel 3370. 


85 


86 Gard. Bull. Sing. 38(1) (1985) 


1-1.2 X 0.5-0.6 mm, broadly rounded at apex; anthers 4-8, septate (at least before 
full maturity), c. 0.8-1.5 mm long, connate (without free apices); androphore 
rather broad, c. 0.1-0.4 mm long. Female perianths ellipsoid to broadly ovoid, 
stout, + coriaceous, c. 3.5-5 * 3-4.5 mm, split at anthesis to c. 14, valves 0.7-1 mm 
thick; pedicels stout, 1.5-5.5 mm long; ovary broadly ovoid-ellipsoid, c. 2.5-3 x 2.5 
mm, glabrous, stigma sessile, not-lobed, c. 0.1 x 0.5 mm. Fruits 2-10 per in- 
fructescence, ellipsoid, at base and apex rounded or sometimes subacutish after 
drying, 3.4-5.5 x 2.5-3.5 cm, glabrous, drying dark brown, without or with few 


coarse tubercles, dry valves 2-4(-5) mm thick; stalk 5-13 mm; perianth not persis- 
TEN. 


Distribution. Moluccas (Morotai to Kai Isls.), Aru Isls., New Guinea (not in 


Morobe and Milne Bay Prov. of Papua New Guinea), not in Bismarck Arch. and 
Solomon Isls. 


MOLUCCAS: b.b. s.n. (April 1920), 16468, 23187, 23749, 24864, 24879, 25176, 25825, (Buwalda 
636) 25981; Beguin 1400; Buwalda 5628, 6112; Idjen & Mochtar 362; Jaheri 710; Kostermans (b.b. 
33725) 13, (b.b. 33921) 251, 767 (p.p.), 1123, 1685; Kuswata & Soepadmo 3; Lam 3463, 3538; Pleyte 
379; Robinson 235; de Vogel 3069, 3094, 3114, 3370, 3491, 3713, 3765, 3815, 3857, 3930, 4164, 4518; 
Teysmann & de Vriese s.n.; de Vriese s.n. 


ARU Isls.: (Buwalda 249) b.b. 25282; Buwalda 4994. 


IRIAN JAYA (incl. Vogelkop): Aet & Idjan 487; b.b. 15901, 22268, 22532, 32682; BW 24, 343, 498, 
533, 536, 1151, 1280, 1366, 1433, 1733, 1754, 1766, 1812, 1838, 1839, 2131, 2207, 2208, 2209, 2210, 2211, 
2227, 2428, 2535, 2687, 2699, 2948, 2950, 3985, 4089, 4369, 4446, 4460, 5191, 5413, 5844, 5923, 6002, 


6539, 6964, 7404, 7689, 7836, 8174, 9194, 9868, 10160, 10835, 10870, 11844, 11869, 11900, 11905, 12409, » 


12990, 13032, 13795, 14929, 15628, 15648, 15653; Gjellerup 180, 180b, 407; Kostermans (b.b. 33355) 


128, (b.b. 33414) 201; (b.b. 33664) 2652, 2669; Kostermans & Soegeng 103, 485; Pleyte 686; van Royen ° 


4510; van Roygen & Sleumer 6193, 7051; Soehoed 31; Zippelius s.n. (180) — Biak & Japen Isl.: Aet & 
Idjan 396, 749; b.b. 1009, 30272, 30393, 30570. 


PAPUA NEW GUINEA: Baldwin UPNG 5752; Brass 7068; Craven & Schodde 739; Darbyshire 908; 
Hart 5007; Hoogland 5021; Jacobs 9052; LAE 73978; NGF 7153, 8214, 10353, 13267, 27471, 34366, 
35623, 36015, 48157, 48467; Pullen 8181; Saunders 920, 1104; Schodde & Craven 4289, 4470; Schodde 
4506; Womersley 3694, 3743. 


Cultivated: (Singapore) Cantley s.n. (1886), Ridley 186, (Furtado) SING 34863 — (Sumatra) Forbes 
1155 A; Teysmann s.n. — (Java) Forbes 1218 c, e; Rastini 90; Sutrisno 66. 


Ecology. Primary and secondary forest, on alluvial soils (sandy and clayey soils), 
especially common in the coastal plains of Vogelkop; also in swampy forest (with 
Pometia), in forest inundated by heavy rains or in stagnant water; also on well- 
drained porous volcanic soils, or close to limestone outcrops, in ridge forest, or in 
Castanopsis forest (at 530 m., in Vogelkop); 0-700 m alt. Sometimes recorded as a 
solitary emergent tree. Flowers and fruits throughout the year. The trees may bear 
flowers and fruits simultaneously.. 


Vernacular names. See Sinclair, p. 147. 


Uses. The wood is heavy, easily worked (easy to cut), not very durable. Fruits 
taste sour, and are edible (Sepik Dist.). The fruit wall is used in rodjak, and in 
manisan (a sweet pickle) (Moluccas). Extract of bark is used as a drug against 
‘““nenyakit keputihan’’ by expecting women, also against hepatitis (Moluccas). The 
fruits are gathered and eaten by the Gogodala tribe, Western Distr., Papua; once 
reported as found planted at a former village site. Fruits eaten by birds (e.g., 
pigeons, parrots), apparently swallowed whole. 


——.. J 


New account of Horsfieldia 2 87 


The trees are several times recorded as beautiful, and recommended as an 
ornamental. 


The many vernacular names indicate that the tree is widely known by local 
people. 


NOTES 


1. Fieldnotes. Recorded as a striking tree, with pendulous branches (twigs up to 
2m), and leaves drooping and distichous (resembling compound pinnate leaves), 
glossy above. Bole often mentioned as very straight, in old specimens with rotten 
core. Recorded as with or without buttresses; these low or up toc. 1.5(-2.5) m high, 
up to 1.5 m out, up to 8(-20) cm thick, sometimes recorded as having small stilt 
roots. Bark brownish, smooth or usually recorded as shallowly fissured, or often as 
slightly to strongly peeling off in small scales; exudate pale red-brown, watery. 
Sapwood colour usually pale yellowish or straw, usually gradually passing into the 
slightly darker or reddish-cream heartwood. Fruits pinkish, pink-orange, pink- 
brown, orange, red-brown, or deep red; aril orange-red or bright red. Flowers 
bright yellow, dark yellow, or orange-yellow, slightly fragrant or not; pollen pale 
yellow or whitish. 


2. Variation and resembling species. Horsfieldia sylvestris is a homogenous 
species; it only rather varies in the hairiness of twigs, leaf-bud, and (juvenile) 
leaves. Very hairy specimens were described as var. villosa by Warburg, but many 
forms intermediate to the typical occur. Wildenow’s H. salicifolia has the leaves 
woolly beneath. A comparatively slightly hairy (i.e., short-haired) specimen is, 
e.g., BW 10835, from the vicinity of Manokwari (Vogelkop); very much hairy 
forms are, e.g., BW 536, 2535, 7836, 11905, 13795, b.b. 32682, or Kostermans 128 
(b.b. 33355), material which partly came from the same area. 


Sterile specimens may be confused with certain forms of H. hellwigii, sometimes 
with similarly hairy twigs and almost similarly long leaf-buds. In that species the 
leaves are usually narrower, the perianth subglobose, not clavate, and the fruits 
smaller and hairy; in the present species the fruits are generally larger, and always 
glabrous. 


In H. sylvestris the number of anthers constituting the elongate androecium is 
rather difficult to ascertain; I counted 4-8 (8-16 thecae), Sinclair had it as 8-10. 


3. Warburg placed H. sylvestris and H. ralunensis in his sect. Orthanthera, which 
also included H. iryaghedhi from Ceylon, because all three species have rather 
elongated and angular male perianths clustered into flower heads. The first two 
named species, however, have the flowers only clustered in immature infloresc- 
ences, and are here not considered closely related to H. iryaghedhi on various 
grounds. 


20. Horsfieldia australiana S.T. Blake Fig. 1A(20); 12 


Horsfieldia australiana $.T. Blake, Austr. J. Bot. 2 (1954) 124, Pl. 5; Sinclair, Gard. Bull. Sing. 28 
(1975) 6, pp., for the Australian specimens only. — Type: S.7. Blake (BRI; iso: SING, n.v.). 


Tree 6-25 m. Twigs terete or lined or ridged towards the top 2-5(-12) mm diam., 
early glabrescent, tomentum pale brown to grey-brown, with minute hairs 0.1 mm 


Fig. 12. 


88 


Horsfieldia australiana §.T. Blake 

a, twig portion with male inflorescences, twig not ridged, x 12; b, ditto, twig lined or ridged, 
x 42; c, opened male flower, showing androecium and schematic longitudinal section of 
androecium, x 6; d, ditto of different specimen, mature male flower closed, opened, and 
schematic longitudinal section of androecium, 6; e, portion of twig with female inflor- 
escences axillary to fallen leaves, x 2; f, mature female flower and longitudinal section of the 
same, note pubescence at base of ovary, X 6; g, portion of twig with infructescence, x ¥2.—a 
& c, from Dunlop 3585; b & d, from Hyland 2724; e & f, from Smith 11913; g, from Hyland 


2d51; 


New account of Horsfieldia 2 89 


or less; bark faintly striate, when older not flaking, lenticels usually conspicuous. 
Leaves in 2 rows, membranous or chartaceous, elliptic oblong to lanceolate, 
broadest at about the middle, or more or less parallel-sided and broadest below or 
above the middle, 10-24 x 3-7 cm, base attenuate, tip obtusish or acutish to 
acute-acuminate, often with bluntish tip; upper surface drying dull olivaceous to 
bright brown, not finely pale-pustulate, lower surface early glabrescent, hairs 
minute 0.1 mm or less, the midrib often rather reddish-brown and usually remain- 
ing sparingly minutely scaly-stellate hairy, lower surface without brown-black dots; 
midrib flattish to slightly raised above; nerves 10-17 pairs, above thin, sunken or 
flat, or slightly raised, beneath with the submarginal arches rather regularly looping 
but inconspicuous; tertiary veins forming a lax network, inconspicuous; petiole 3-7 
x 1.5-2 mm, leaf bud 8-15 x 1-2 mm, with hairs c. 0.1 mm. Inflorescences sparsely 
to densely pubescent with hairs 0.1-0.2 mm long hairs, in C&’ and Q: 2 or 3 times 
ramified, (1.5-) 3-8 x 2-5 cm, common peduncle 0.5-1.0 cm long; bracts elliptic to 
broadly triangular, acutish, 2-6 mm long, short-pubescent, caducous. Flowers in 
loose clusters of 2-6 (in Q: 1-3) each; perianths 2-valved, rather sparsely pubescent 
with hairs 0.1 mm or less long, pedicels slender, thinly pubescent, at base inarticu- 
late. Male perianth ellipsoid or subglobose, slightly laterally compressed, (2-)2.5- 
3.3 X (1.5-) 2-3.0 mm, above and below rounded, pedicel not tapering, 1-2 mm 
long; perianth at anthesis cleft to c. ¥2-way, valves c. 0.2 mm thick. Androecium 
moderately laterally compressed, above subtruncate to broadly rounded, below 
somewhat attenuate, (1.8-) 2.0 x 1.5-2.0 mm; anthers (10?-) 12-14, almost straight, 
the free portions at apex c. 0.1-0.2 mm long and these slightly or much incurved; 
the column rather wide, spongy, the apical “4-2 broadly hollowed out but the basal 
portion more or less protruding into the upper part of the hollow; androphore 
rather narrow, 0.1-0.2 mm long. Female perianth ellipsoid, c. 2.5(-3) xX 2.0(-2.5) 
mm, cleft at anthesis to c. 2-way, valves c. 0.2 mm thick, pedicel 1-2 mm long; 
ovary ovoid, c. 2.0 X 1.5 mm, towards base with dense tomentum of minute hairs 
0.1 mm or less, style minutely 2-lobed, c. 0.2 mm broad. Fruits 3-8 per infructesc- 
ence, ellipsoid, top rounded or faintly pointed, 1.8-2.2 x 1.1-1.4 cm, with the 
surface granulate and with or without a few coarse tubercles, glabrous except at the 
very base; dry pericarp c. 2 mm thick; stalk 2-4 mm long; perianth not persisting. 


Distribution. Australia: Northern Territory, northern Queensland. 


AUSTRALIA. Northern Territory: Byrnes NB 1259 (NT. 14918); Dunlop 3585; Must & McKean B 
687 — Queensland: Hyland 2551, 2557, 2724, 3123, 5516; Smith 11762, 11913; Stocker 1043. 


Ecology. Riverine forest, gallery forest in gullies of sandstone areas, sheltered 
gorge forest, monsoon forest on sandy soils; 0-200 m alt. Flowers from August to 
October, fruits from September to January. 


NOTES 


1. Fieldnotes. Bark fissured or tessellated, usually flaky, stem often recorded as 
+ fluted, buttressed. Blaze exudate watery, red; blaze smells like ants. Wood 
whitish. Flowers yellow or orange, the female noted as strongly and sweetly scented 
(as the flowers of Alocasia macrorhiza or the fruits of Passiflora edulis); the males 
recorded as scentless. 


90 Gard. Bull. Sing. 38(1) (1985) 


2. Possibly two forms can be recognized, but the material at hand prevents a final 
decision. Specimens from Northern Territory, incl. the type and Must & McKean B 
687 (C'), Dunlop 3585 (CC), Byrnes NB 1259 (fr.), have the twigs not or only 
indistinctly ridged, the leaf blades rather elliptic-oblong (not oblong-lanceolate), 
possibly relatively longer petioles, the blades more of a membranous texture with 
+ prominent lateral nerves, and possibly the male perianth broader, i.e., broadly 
ellipsoid or subglobose. Specimens from Queensland have usually rather leathery 
leaves, usually with flattened or sunken lateral nerves, the blades usually more 
oblong-lanceolate, with shorter and broader petioles, and the twigs apparently 
more distinctly ridged or even winged. However, specimens which distract from 
this image exist, e.g., the sterile specimen Hyland 3123 from Northern Terr. has an 
‘easternly’ habit, or the sterile specimen Smith 11762, from Queensland, which has 
the leaves rather membranous and nerves prominent. 


3. Sinclair accepted the present species as including specimens from New Guinea 
which are presently regarded as representing a separate new species, H. sinclairii. 


21. Horsfieldia crux-melitensis Markgraf Fig. 1A(21); 13 a-c 


Horsfieldia crux-melitensis Markgraf, Bot. Jahrb. 67, 2 (1935) 148; Sinclair, Gard. Bull. Sing. 28 (1975) 
26 (cruxmilitensis) — Type: Schlechter 19246 (B, n.v.; iso: K, L; E, G, NY, S, Z, n.v.). 


Shrub or treelet 2-4 m. Twigs terete, not ridged, towards the apex 1.5-3(-4) mm 
diam., glabrescent, tomentum rusty, of hairs c. 0.1 mm; bark finely striate, when 
older not flaking, lenticels absent or inconspicuous. Leaves in 2 rows, membra- 
nous, elliptic to obovate-oblong, broadest usually above the middle, 12-27 x 
5.5-11.5 cm, base attenuate, tip acute-acuminate; upper surface drying dark 
brown, without or with very minute paler dots, lower surface with persistent 
tomentum of rather sparse hairs (c. 0.1 mm) especially on the midrib, or late 
glabrescent, without dark dots, the nerves not contrasting in colour; midrib above 
slightly raised; nerves 10-15 pairs, sometimes with additional intercalary nerves, 
thin and flat above, much raised beneath, the submarginal arches distinct but not 
very regularly shaped; tertiary veins forming a lax, rather indistinct network; 
petiole 10-16 X 1.5-2.5 mm; leaf bud 7-12 x 1-2 mm, with hairs 0.1-0.2 mm. 
Inflorescences situated in between the leaves, woolly-pubescent with stellate- 
dendroid hairs 0.1-0.2 mm, 2(-3) times ramified, in CG’ and 9: rather few-flowered, 
c. 1.5-5 xX 1.5-4 cm, common peduncle 0).6-1.5 cm long; bracts c. 0.5 mm, pubes- 
cent, early caducous. Flowers solitary or up to 3 together; perianths 2-valved, with 
scattered stellate hairs c. 0.1(-0.2) mm, densest towards base, pedicels tapering, 
pubescent, at base inarticulate. Male perianth subglobose, gradually passing into 
the strongly thickened and tapered pedicel, together forming the long-clubshaped 
flower c. 9-11 xX 2-3.2 mm; perianth broadly rounded above, somewhat laterally 
compressed, c. 2-3 X 2-3.2 mm, pedicel c. 7-8 X 2-3 mm, + obconical, rather 
densely pubescent; perianth at anthesis split to c. 4%-'%, i.e., to 0.3-0.6 mm deep, 
valves c. 0.2-0.3 mm thick, the perianth-wall lower down c. 0.6-0.8 mm thick. 
Androecium a club-shaped body, the top + rounded, c. 1.5-2.5 x 0.7-1.2 mm, 
with 3-5 stellately arranged, connate anthers at the apex, c. 0.2-0.3 mm long, 
androphore thick, subcylindrical, the surface + wrinkled-bullate and more striate 
towards the base, glabrous; central column up to the apex solid. Female perianth 
and pedicel together forming an obconical flower similar to O’ flowers, c. 8 xX 3 


Fig. 13. 


Horsfieldia crux-melitensis Markgraf: a, leaf, x ‘2; b, opened male flower, showing club- 
shaped androecium, X 6; c, female flower, opened, showing pubescent ovary with minute 
narrow bilobed style, x 6. — H. clavata de Wilde: d, portion of twig with infructescence with 
mature fruit, x 2; e, mature male flower, lateral view, x 6; f, ditto, opened, showing 
club-shaped androecium, x 6. — H. squamulosa de Wilde: g, habit of leafy twig with male 
inflorescences. —a & c, from LAE 73830; b, from Schlechter 19246 (type); d, from Hoogland 
3623; e & f, from Hoogland 3663 (type); g, from Brass 7221. 


91 


92 Gard. Bull. Sing. 38(1) (1985) 


mm; perianth c. 2.2 x 3 mm, split at anthesis for c. 0.2-0.3 m deep; pedicel broadly 
obconical, c. 6 X 2.6 mm; ovary obvoid, c. 2 x 1.5 mm, densely pubescent through 
hairs c. 0.1 mm long or less, style together with the 2-lobed stigma c. 0.2-0.3 mm . 
long. Fruits (according to young infructescences in LAE 73830) 1-4 each, possibly 
becoming more than 1.6 cm long, pubescent; stalk much elongated, and thickened 
towards the fruit, 10-14 mm long; perianth not persisting. 


Distribution. Papua New Guinea: Morobe Prov. 


PAPUA NEW GUINEA. Morobe Prov.: LAE 73822, 73830; NGF. 24092; Rau 550; Schlechter 
19246. 


Ecology. Mixed rain forest, lowland forest, common in wet areas; 0-50 m. 
Flowers from January to May; fruits in March. 


NOTES 


1. Fieldnotes. Shrub or treelet, 2-4 m; bark grey or dark green, wood cream or 
white. Flowers cream or orange. Fruits yellowish or red. 


2. Fruits ripening red are said to have been mentioned on the label of LAE 
73530; in L only female flowers and young fruits are present. Mature fruits have not 
been seen by me. 


3. Specimens from the Northern Province, presently assigned to a new species H. 
clavata, were included by Sinclair in the present species. 


4. | have not seen the duplicates of the para-type, Schlechter 17408 (G, NY). 
According to Sinclair’s notes on p. 27, its leaves are rather narrowly elliptic or 
elliptic or elliptic, not broadly elliptic. This means the specimen could belong to different 
related species, e.g., H. clavata, or rather H. squamulosa, which is also known 
from the Morobe Province. 


22. Horsfieldia clavata de Wilde, sp. nov. Fig. 1A(22); 13 d-f 


Horsfieldiae crux-melitensis atque eodem modo Horsfieldiae squamulosae affinis quoad androecium 
clavatum, sed differt a H. squamulosa floribus masculis multo minoribus, a H. crux-melitensis pedicellis 
attenuatis. — Type: Hoogland 3663 (L; iso: BM, K; CANB, n.v.). 


Shrub or tree, 3-6 m. Twigs terete, not ridged, 1.5-3 mm diam., glabrescent, 
tomentum grey-rusty, of hairs c. 0.1 mm long; bark finely striate, when older not 
flaking, lenticels absent or inconspicuous. Leaves in 2 rows, membranous, elliptic 
to oblong, broadest at or somewhat above the middle, 7-18 x 3-6 cm, base short- to 
long-attenuate, tip acute-acuminate (in Hoogland 3523 c. 2 cm, caudate); upper 
surface drying olivaceous, without minute paler dots, lower surface with persistent 
tomentum of rather scattered stellate-dendroid, scale-like hairs c. 0.1-0.2 mm 
especially on midrib, without larger dark brown dots, the nerves not much contrast- 
ing; midrib slender above, raised; nerves 10-20 pairs (including some weaker 
intercalary nerves), above thin and flat or slightly raised, beneath much raised, with 
distinct, rather regularly looping submarginal arches; tertiary veins forming a lax, 
rather indistinct network; petiole 7-14 x 1-1.5 mm; leaf bud 7-10 x 1-1.5 mm, with 
hairs c. 0.1 mm. Inflorescences situated in between the leaves, delicate 1-2(-3) 
times ramified, lowest side branch from near the base, in CO’ rather many-flowered, 


New account of Horsfieldia 2 93 


2-3 x 1.5-2 cm, in Q: 2-3-flowered, 1-2 cm long; axes finely scaly-pubescent with 
hairs c. 0.1 mm or less; bracts densely pubescent, c. 1-1.5 mm long, caducous. 
Flowers solitary or 2-3 together; perianths 2-valved, rather densely pubescent with 
stellate-dendroid hairs c. 0.1 mm; pedicels at base inarticulate. Male perianth 
subglobose, gradually passing into the strongly tapering pedicel, together forming a 
club-shaped flower c. 4-5.5 xX 1.5-2.2 mm; perianth rounded above, c. 1.5-2 xX 
1.5-2.2 mm, pedicel c. 2.5-3.5 x 1-1.5 mm, rather densely pubescent; perianth at 
anthesis split to c. Mio (i.e., for c. 0.2 mm only), valves c. 0.2 mm thick, the 
perianth-wall lower down c. 0.5-0.7 mm thick. Androecium a club-shaped body c. 
1.5 x 0.7 mm, at apex consisting of 3 anthers (or c. 6 thecae;; c. 0.3 mm long, + 
arranged into a star shape, without free apices; the androphore thickish sub- 
cylindrical, slightly bullate-striate, glabrous; central column not hollowed out at 
apex. Female perianth ellipsoid, c. 1.8(-2) x 1.2 mm, split at anthesis to c. 4%, 
valves c. 0.2 mm thick; pedicel + slender, c. 2 mm long; ovary ovoid, c. 1.0 x 0.6 
mm, densely pubescent with stellate-scaly hairs c. 0.1 mm or less, style c. 0.4 mm 
long, stigma 2-lobed, c. 0.2 mm long. Fruits 1 (or 2) per infructescence, broadly 
ellipsoid-ovoid, base broadly rounded, top + acuminate, c. 2 mm beaked, excl. the 
1.5-2 mm long pseudo-stalk c. 1.3 X 1.0 cm, pubescent with hairs c. 0.1 mm long, 
drying brownish, without lenticels; dry valves c. 1 mm thick; seed ellipsoid; stalk 
6-10 mm long; perianth not persistent. 


Distribution. Papua New Guinea: Northern Prov. 


PAPUA NEW GUINEA. Northern Prov.: Hoogland 3523, 3623, 3663. 


Ecology. Locally common in regrowth in tall lowland forest, on well-drained soil; 
0-50 m. Flowers and fruits in August. 


Vernacular name. Hamana (Orokaiva lang., Mumuni). 
NOTES 
1. Fieldnotes. Shrub or treelet; flowers yellow; fruits orange or red, aril red. 


2. Related to H. squamulosa and H. crux-melitensis which have a similar club- 
shaped androecium. H. squamulosa differs by its slender, male pedicels. The 
pedicel, and hence the whole male flower of H. crux-melitensis is similarly club- 
shaped as in the present species, but about twice as large; its leaves are also larger, 
drying to a darker colour. In H. crux-melitensis both male and female flowers have 
much thickened pedicels. 


3. Sinclair included the specimens of the present species in H. crux-melitensis. 


23. Horsfieldia squamulosa de Wilde, sp. nov. Fig. 1A(23); 13 g 


Horsfieldia species androecio clavato, H. crux-melitensis atque eodem mode H. clavatae affinis, 
differt pedicellis gracilibus non-attenuatis — Type: Henty & Barlow NGF 42995 (L; iso: K; A, BRI, 
CANB, n.v.). 


Shrub or tree, 1-10 m. Twigs terete, not ridged, towards the apex 1-3 mm diam., 
glabrescent, tomentum of minute rust-coloured hairs 0.1-0.2 mm long; bark finely 
striate, when older not flaking, lenticels fine and inconspicuous. Leaves in 2 rows, 


94 Gard. Bull. Sing. 38(1) (1985) 


membranous, elliptic to lanceolate, broadest at or above the middle or + parallel- 
sided, 4-20 x 0.7-5 cm, base attenuate, tip up to 3.5 cm long, acute-acuminate, 
upper surface drying dark brown, with or without very minute paler dots, lower 
surface with persistent tomentum (especially on the midrib) or glabrescent, hairs 
0.1-0.2 mm long or less, without larger dark brown dots, the nerves not contrasting 
in colour; midrib slender above and slightly raised; nerves 10-25 pairs, including 
some weaker intercalary ones, thin and flat above, much raised beneath, with the 
submarginal arches regularly looping or not, distinct or not; tertiary veins forming a 
lax network, distinct or indistinct; petiole 6-13 x 1-1.5 mm; leaf bud c. 10 xk 1 mm 
with dark rust-coloured hairs 0.1-0.2 mm long. Inflorescences in between the 
leaves, 2(-3) times ramified (lowest branch 0-5 mm from the base), few to rather 
many-flowered, in G&: c. 1.5-3 X 1-2 cm, in 9 few-flowered, c. 1-3 cm long; axes 
woolly pubescent through stellate scaly or dendroid hairs 0.1-0.2 mm long; bracts 
pubescent, 0.5-1.5 mm long, caducous. Flowers 1-3 together; perianths 2-valved, 
stellate-scaly hairs scattered, densest towards the base; pedicel + densely scaly- 
pubescent, at base inarticulate. Male perianth ellipsoid to broadly obovoid, 2.0-3.0. 
x 1.5-2.2(-2.4) mm, upper part rounded, at base long- to short-attenuate; pedicel 
2-3.5(-6) mm, slender; perianth at anthesis split to c. “%-% (c. 0.1-0.4 mm deep 
only), valves 0.1-0.2 mm thick, the perianth-wall lower down 0.5-0.8 mm thick. 
Androecium a cylindrical club-shaped body 1.5-2.5 x 0.5-0.8 mm, at apex consist- 
ing of 3-4 anthers (or 6-8 thecae) 0.3-0.5 mm (the anthers + stellately arranged, 
with free apices 0-0.2 mm), and lower down a thickish sub-cylindrical androphore, 
in the upper '3-%4 wrinkled-bullate or warted, in the lower 2-74 either minutely 
scaly-hairy or surface striate but glabrous; central column not hollowed out at 
apex. Female perianth broadly ellipsoid, 2-2.5 x 1.5-2.1 mm, split at anthesis to c. 
4-3, valves c. 0.2(-0.3) mm thick; pedicel 2-3.5 mm long; ovary ovoid, c. 1.5 X 
1.2-1.4 mm, densely pubescent through stellate-scaly hairs c. 0.1 mm or less, style 
c. 0.2 mm long, stigma distinctly 2-lobed, 0.2-0.3 mm long. Fruits 1-3 per in- 
fructescence, broadly ellipsoid-ovoid, base rounded, top rounded to acutish, ros- 
trum if present 1-3 mm, excl. the 1.5-5 mm long pseudostalk c. 1.0-1.6  0.7-1.1 
cm, sparsely to densely pubescent through hairs c. 0.1 mm long, drying blackish, 
without lenticels; dry valves c. 0.5(-1.0) mm thick; seed ellipsoid; stalk 10-12 mm; 
perianth not persisting. 


Distribution. Papua New Guinea: Morobe Prov., Northern Prov., Milne Bay 
Prov. (incl. Normanby Isl.), Western Prov. 


PAPUA NEW GUINEA: Brass 7221; Carr 16192; LAE 67148, 70239, 71160; NGF 23574, 28894, 
31751, 31888, 34088, 38093,.42995, 46892; Pullen 8287; Schlechter 17408 (n.v.) 


Ecology. Scattered or locally plentiful shrub or small understorey tree, 1-6(-10) 
m in rain forest; on slopes and ridges, creek banks, edges of (sago) swamp forest. In 
Normanby Isl. in Eucalyptopsis-dominated forest, in New Guinea found associated 
in forest with dominant Castanopsis, or with Lithocarpus, Anisoptera and Hopea- 
dominant; 0-500 m. alt. Flowers and fruits throughout the year. 


NOTES 


1. Fieldnotes. Shrubs or treelets, recorded with the branches + whorled, or 
horizontal. Bark smooth, grey-brown; red exudate; wood cream to straw. Flowers 
yellow. Ripe fruit orange to orange-red, aril bright red. 


New account of Horsfieldia 2 95 


2. By the similar shape of the androecium closely related with H. clavata and H. 
crux-melitensis. Most collections were acquired after Sinclair's time. Only one 
specimen was included by Sinclair in H. subtilis var. schlechteri, which is presently 
again accepted as a separate species, H. schlechteri. 


3. The androphore in Pullen 8287, from Milne Bay Prov., is glabrous, whereas 
those of the type specimen, and other specimens from Western Prov. are densely 
minutely scaly-hairy on the lower part of the androecium. It is possible that this 
difference has a taxonomical significance, but more material is needed to decide on 
it. 


24. Horsfieldia ampla Markgraf 


Horsfieldia ampla Markgraf, Bot. Jahrb. 67, 2 (1935) 148 — Type: (Sepik) Ledermann 9639 (B, +, n.v.). 


Small tree, 4-5 m. Twigs terete. Leaves large, cuneate-obovate, up to 40 x 16 
cm, base + attenuate, tip short-acuminate; nerves 16-18 pairs, straight, sharply 
raised beneath and connected before the margin. Petiole, 1 cm long. Inflorescences 
on the older wood, large, to 25 cm long and 10 cm wide, glabrescent, loosely 
flowered. Male flowers yellow, clavate, 4 x 2 mm (excl. pedicel?), the perianth 
2-valved, split at anthesis to hardly 4. Staminal column thick; anthers to c. 10, the 
androphore about as long as the anthers or slightly shorter. 


Distribution. NE. New Guinea, Sepik Prov., Aprilfluss. Mountain slope near 
camp 18. Known only from the type. 


Ecology. Dense, very humid forest, on mountain slope at 200-400 m alt. Male 
flowers date 12 November, 1912. 


NOTES 


1. This species is known only from the type. It is keyed out by Markgraf against 
H. crux-melitensis, with which it has in common the clavate flowers. In the key is 
mentioned that the perianth, other than the androecium, is largely hollow. He 
mentions in a note that the species is peculiar amongst the New Guinean Horsfiel- 
dias because of its large male flowers, which in other species are smaller and almost 
always broader than long, and hence that it is without close relatives. 


2.. H. ampla was not mentioned by Sinclair. 


25. Horsfieldia ampliformis de Wilde, sp. nov. Fig. 1B(25); 14 


Horsfieldia species inflorescentiis masculis grandis, c. 30 cm longis, attamen pubescentibus, a H. 
ampla Markgraf differt perianthiis latioribus (c. 3 mm latis) atque antheris 7. — Type: (New Guinea, 
Sepik Dist.) Hoogland & Craven 11085 (oC fl.). (L; iso: K; A, BRI, CANB, LAE, US, n.v.). 


Tree 5-8 m. Twigs stoutish, terete, when young thinly ridged, towards apex 
4-7(-10) mm diam., early or late glabrescent, pubescence + woolly, of hairs c. 
0.2-0.5 mm long; bark coarsely striate, lenticellate, when older not flaking. Leaves 
in 2 rows, thickly membranous, elliptic-oblong to oblong, (18-)25-38 x (6-)7-13 
cm, base short to long-attenuate, tip attenuate-acuminate; upper surface drying 
dark brown, lower surface early or late glabrescent or with subpersistent tomentum 
of rather scattered stellate hairs 0.2-0.5 mm long; larger dark brown dots absent; 


Horsfieldia ampliformis de Wilde. 
a, twig apex with leaves, x '2; b, portion of twig with male inflorescence axillary to fallen leaf, 
x 2; c, mature male flower, perianth opened, showing androecium, X 6; d, portion of twig 
with female inflorescence, x 2; e, female flower, opened, showing finely pubescent ovary 
and minute 2-lipped stigma. — a-c, from Hoogland & Craven 11085; d & e, from Craven & 
Schodde 1463. 


96 


New account of Horsfieldia 2 97 


midrib above + slender, flattish; nerves 18-22 pairs, above thin, flat or sunken; the 
submarginal arches beneath not very prominent; tertiary venation forming a lax 
network, indistinct on both surfaces; petiole short, 4-6 x 3-4 mm; leaf bud 25-40 
mm long, with hairs 0.2-0.5 mm. Inflorescences situated below the leaves, in C: 
large, many-flowered, 4-5 times ramified, c. 25-35 x 20-30 cm, common peduncle 
40-50 mm; in 9: c. 9-10 x 6-8 cm; all branches rather loosely pubescent with hairs 
0.2-0.5 mm long; bracts (only seen in 2) c. 5 mm long, caducous. Flowers 2-5 
together in CO and Q, flowers and pedicels loosely pubescent, hairs (0.1-)0.2-0.3 
mm long, in 9 the perianth glabrescent towards apex; perianths 2-valved; pedicel 
at base inarticulate. Male perianths broadly obovoid, laterally + flattened, c. 3-3.3 
x 3-3.2 mm, at apex obtuse to broadly rounded, at base shortly tapering into 
pedicel 2-4 mm long; perianth largely hollow, at anthesis split to c. %, valves c. 0.3 
mm thick. Androecium small, + flattened, incl. androphore c. 2.5 x 1-1.2 mm, 
broadly rounded at apex; anthers 7, when young indistinctly septate, synandrium 
1.5-1.8 xX 1-1.2 mm, free apices of anthers 0.1-0.2 mm; androphore 0.8-1.0 x 
0.5-0.6 mm; the column at apex narrowly hollowed for c. “4-3. Female perianths 
broadly ovoid, c. 3 X 2.6-2.8 mm, split at anthesis to c. 7, valves c. 0.3 mm thick; 
pedicel 1-2 mm long; ovary broadly ovoid, c. 2-2.2 xX 1.8-2.0 mm, densely pubes- 
cent with hairs 0.1 mm or less, stigma short, not or hardly lobed, c. 0.1 X 0.4 mm. 
Fruits not seen. 


Distribution. Northern Papua New Guinea: Sepik Prov., Morobe Prov. 


PAPUA NEW GUINEA. Craven & Schodde 1463; Hoogiand & Craven 110835. 


Ecology. Lower montane rainforest, 1200-1300 m alt. Flowers in April and 
August. 


Vernacular name. Guma (Sepik Prov., Waskuk lang.). 
NOTES 
1. Fieldnotes. Small tree, c. 8 m high. Flowers medium green, yellow at anthesis. 


2. Close to H. ampla Mkgf., a species of which I have seen no material. 
According to the description it differs by the more elongate, possibly glabrous, 
perianths c. 4 X 2 mm, the androecium with 10 anthers, and the glabrescent 
inflorescences. The present new species has in common with H. ampla the peculiar- 
ly long-stalked androecium and the large male inflorescences. H. ampla was 
collected at 200-400 m alt. 


3. Known from a male and a female flowering specimen. The perianths of the 
female specimen, Craven & Schodde 1463, from Morobe Prov., are glabrescent in 
the upper half. This could also be H. ampla as well. Moreover, as the hairs on the 
leaf bud are slightly shorter than those of the male specimen it is difficult to 
distinguish H. ampliformis from the variable and widespread H. laevigata. 


26. Horsfieldia angularis de Wilde, sp. nov. Fig. 1B(26) 
Ramuli angulares vel biporcati, perianthio masculo subgloboso, 2-3 mm diam., basin versus pubes- 


centi, 2-4-valvato fere usque ad basin, antheris 12-20, erectis, ovario pubescenti, fructibus breviter 
ellipsoideis, 1.7-2.0 cm longis, minute pubescentibus vel glabrescentibus. — Type: Vogelkop Penins., & 


98 Gard. Bull. Sing. 38(1) (1985) 


fl.. BW 5828 (L). 


Tree 15-30 m. Twigs 2-angular from the two ridges from petiole to petiole, lower 
down subterete with two distinct ridges, 3-7(-10) mm diam., early glabrescent, 
tomentum grey-brown, with hairs 0.1 mm or less; bark striate, distinctly coarsely 
lenticellate, when older not flaking. Leaves in 2 rows, membranous to thinly 
chartaceous, oblong to oblong-lanceolate, broadest usually in the middle, 10-27 x 
3-7.5 cm, base attenuate, tip acute-acuminate; upper surface drying pale to dark 
brown, often finely paler pustulate, lower surface glabrescent, hairs very minute 
grey stellate, less than 0.1 mm; without brown or blackish dots; midrib flattish or 
slightly raised above; nerves 12-15 pairs, above thin and flattish or slightly sunken; 
tertiary venation forming a lax network, faint; petiole 7-15 x 2-3 mm; leaf bud 
10-15 x 2-2.5 mm, with hairs 0.1 mm. Inflorescences rather densely pubescent with 
hairs 0.2-0.3 mm; in Gand Q: 2 or 3 times ramified, rather few-flowered, c. 3-4 x 
2-2.5 cm, common peduncle 3-6 mm; bracts not seen, caducous. Flowers (in GC) 
generally 2-4 together; perianths in CO’: 2-4-, in Q: 2(-3)-valved, pubescent in the 
lower half with hairs 0.1(-0.2) mm long; pedicels pubescent, at base inarticulate. 
Male perianth in lateral view circular or slightly transversely ellipsoid, slightly or 
not laterally compressed, not collapsing on drying, 1.7-2.3 x 2.2-3.2 mm, above 
and at base (broadly) rounded, pedicel not tapering, 1-2 mm long; perianth at 
anthesis cleft to the base (c. 10), valves (0.2-) 0.3 mm thick. Androecium slightly 
laterally flattened (and in 3- or 4-valved flowers + 3- or 4-angular in transverse 
section), above broadly rounded, 1.2-1.5 xX 1.5-2.2 mm, anthers 12 (2-valved) to c. 
20 (in 4 valved flowers), + erect, not septate, free portions at apex up to 0.1 mm, 
central column at apex narrowly hollowed for (4%-)'2; androphore absent, the 
androecium + broadly attached. Female perianth depressed globose, c. 2.5 X 3-3.2 
mm, cleft at anthesis to c. 4, valves 0.8-1.0 mm thick, pedicel 1-1.5 mm long; ovary 
+ depressed globose-ovoid, c. 1.2 X 1.5 mm, densely short-pubescent, style and 
stigma minute, minutely 2-lobed, c. 0.1 X 0.3 mm. Fruits 5-10 per infructescence, 
short-ellipsoid, 1.7-2.0 x 1.4-1.7 cm, pubescent at very base, with coarse paler- 
coloured lenticel-like tubercles; dry valves thick-woody, c. 3-5 mm thick; seed 
ellipsoid, stalk 3-5 mm long; perianth not persistent. 


Distribution. New Guinea: Vogelkop Peninsula, subdist. Manokwari. 


NEW GUINEA. Irian Jaya, Vogelkop: BW 2340, 5828, 10922, 15752; Kostermans 2635; van Royen & 
Sleumer 6813 — Jayapura Dist. (doubtful): b.b. 31124; BW 5340. 


Ecology. Primary forest; on clayey soils, locally common; 0-600 m alt. Flowers in 
February and August, fruits in February and October. 


Vernacular names. Babijag (Karoon lang.), Bepoes (Hattam lang.), Betelohoi 
and Sebohonggwa (Manikiong lang.). 


NOTES 


1. Fieldnotes. Locally common in primary forest on the coastal plain, up to 600 
m. in Kebar valley. Sometimes buttresses to c. 1 x ¥2 m. Bark sometimes fissured, 
or peeling off in small scales; with red exudate. Sapwood pale brown or white, 
heartwood not discernable or pinkish. Flowers greenish. Fruits yellow-brown or 
yellow, aril orange or red; fruit recorded as sour and edible. 


2. The present new species is much related to H. basifissa of which the sterile 
specimens (presumed to belong to it) are difficult to identify since their twigs are 
rather ridged. H. angularis is distinguished from H. basifissa by (1) the more 


New account of Horsfieldia 2 99° 


strongly ridged and somewhat stouter twigs, (2) the more hairy and variably 
2-4-valved flowers with thicker valves, (3) the hairy ovary, and the thinly pubescent 
ellipsoid fruits; the two species have in common thickish subglobose male (flower) 
perianths, which do not or hardly collapse on drying, and which at anthesis are cleft 
to the base. 


3. Sinclair identified specimens as H. polyantha, a name presently considered a 
synonym of H. laevigata; specimens of the resembling H. basifissa were also 
identified by Sinclair as H. polyantha. 


4. Two sterile specimens, with ridged twigs, b.b. 31/124 and Karstel BW 5340, 
from Jayapura Dist., Irian Jaya, probably belong to the present species. 


27. Horsfieldia iriana de Wilde, sp. nov. Fig. 1B(27) 


Horsfieldia novo-guineensis Warb., Mon. Myrist. (1897) 271, p.p., only the type of H. iriana, not the 
lectotype. 


Myristica nesophila auct. non. Miq., Ann. Mus. Bot. I (1864) 206: Mig., Ann. Mus. Bot. II (1865) 49, 
p.p., as based on Zipelius (139 d), not the lectotype of Myristica aruana Bl. = H. aruana. 


Ramuli angulati vel sulcati. Perianthium in fl. & valvis 2 instructum. Androecium lateraliter com- 
pressum. Antherae erectae. Cum H. aruana comparibilis, sed differt pedicellis perianthio longioribus 
perianthoque sub anthesi usque ad basin fissum. — Type: Zipelius (139 d) (L; iso: K). 


Tree c. 10(?) m. Twigs 2-angular from the two ridges between the petioles, lower 
down subterete but ridged, 3-5 mm diam., early glabrescent, tomentum grey-rusty, 
composed of hairs c. 0.2 mm long; bark rather smooth, when older not flaking; 
lenticels small, distinct. Leaves in two rows, thinly coriaceous, oblong-lanceolate, 
broadest at or above the middle, 17-28 x 4.5-8 cm, base long-attenuate, tip 
acute-acuminate; upper surface drying olivaceous, lower surface early glabrescent 
from stellate-scaly hairs 0.1-0.2 mm long, larger brown-blackish dots absent; 
midrib rather broad and flat above; nerves 13-16 pairs, thin and flat above; tertiary 
veins forming a lax network, slightly raised above but indistinct; petioles 6-11 x 
2-3.5 mm; leaf bud slender, 10-15 xX 1.5-2 mm, with hairs 0.1-0.2 mm long. 
Inflorescences in CG: axillary to the lower leaves, sparsely pubescent with stellate 
hairs 0.2(-0.3) mm long, (2 or) 3(or 4) times ramified, rather many-flowered, 6-8 x 
4-6 cm, common peduncle 5-10 mm long; bracts not seen, caducous. Flowers in 
loose clusters of 2-5, perianth 2-valved, largely glabrous but with some minute hairs 
towards the base; pedicels slender, sparsely pubescent, at base inarticulate. Male 
perianth in lateral view circular to slightly longitudinally ellipsoid, not or slightly 
laterally compressed, blackish brown, not collapsed on drying, 2.3-2.8 x 2.2-2.8 
mm, subacute or narrowly rounded at the top, at base rounded, pedicel not 
tapered, slender, c. 2.5-3 mm long; perianth at anthesis cleft to +4-% deep, valves c. 
0.2 mm thick. Androecium laterally compressed (flattened), c. 1.5 x 1.5-2 mm, 
(broadly) rounded above; anthers c. 10-14, erect, finely septate when young, free 
portions at apex 0.1(-0.2) mm long, column at apex narrowly hollowed for c. 4; 
androphore absent. Female flowers and fruits not seen. 


Distribution. SW. New Guinea (Irian Jaya), known only from the type. 


100 Gard. Bull. Sing. 38(1) (1985) 


Ecology. Nothing known; likely from coastal lowland forest. 
NOTES 


J. Only known from the type specimen, Zipelius (139 d) in L (K, iso). This 
specimen is part of the heterogenous Zipelius material which served for the 
description of Myristica aruana Blume and Horsfieldia novo-quineensis Warb., of 
which the typification is explained under Horsfieldia aruana. The syntype- 
specimens Zipelius (139d) were erroneously included by Sinclair in H. polyantha 
Warb. as accepted by him. This name is here considered a synonym of H. laevigata. 


28. Horsfieldia aruana (B1.) de Wilde, comb. nov. Fig. 1B(28) 


Myristica aruana B\., Rumphia 1 (1837) 191; Sinclair, Gard. Bull. Sing. 28 (1975), 112, 118, 119 122-124, 
in the synonymy of Horsfieldia spicata — Palala aruana Rumph., Herb.Amb. 7(Auct.)(1755), t. 24 — 
H. novo-guineensis Warburg, Mon. Myrist. (1897) 271, t. 23 fig. 1-3, p.p., for the lectotype only. — 
Lectotype: those specimens of Zipelius s.n. at L, annotated by Blume. 


Tree c. 15m. Twigs 2-angular from the two ridges between the petioles, lower 
down subterete though also provided with 2 ridges, 3-5 mm diam., early glabres- 
cent, tomentum minute, of hairs c. 0.1 mm long; bark striate, when older not 
flaking; lenticels small, inconspicuous. Leaves in two rows, membranous, elliptic- 
oblong, broadest at or above the middle, 15-29 x 5-9.5 cm, base attenuate, tip 
acute-acuminate, upper surface drying olivaceous to brown, lower (surface) early 
glabrescent, without larger brown-blackish dots; midrib flattish or slightly raised 
above; nerves 13-15 pairs, slender, flattish; tertiary veins forming a lax network, 
indistinct; petioles 10-15 x 1.5-2.5 mm, leaf bud c. 10 x 1.5 mm, with hairs c. 0.1 
mm long. Inflorescences situated in between or below the leaves, sparsely pubes- 
cent by hairs c. 0.1 mm or less, in C&: 3 or 4 times ramified, 5-8 x 4-5 cm, rather 
many-flowered, common peduncle 5-15 mm long; bracts not seen, caducous. 
Flowers in loose clusters of 2-5 each, perianth 2-valved, glabrous; pedicels slender, 
sparsely pubescent, inarticulated at base. Male perianth in lateral view circular to 
somewhat transversely ellipsoid, laterally compressed, blackish and collapsing on 
drying, 1.5-2 xX 2-2.5 mm, top broadly rounded, base + rounded, pedicel not 
tapering, slender, 1-1.5 mm long; perianth at anthesis cleft to c. 74-%4, valves c. 0.2 
mm thick. Androecium much laterally compressed, c. 1.5 X 2.0 mm, above broadly 
truncate-rounded; anthers (c.) 14-18, finely septate when young, distal free por- 
tions 0-0.1 mm, anther column completely solid or almost so (see notes); 
androphore absent or up to 0.1 (-0.2) mm. Female flowers and fruits not seen. 


Distribution. SW. New Guinea (Irian Jaya); possibly also Aru and Tanimbar Isls. 
(see notes). 


TANIMBAR ISLS.: b.b. 24414 (doubtful). 
ARU ISLS.: Buwalda 4969 (doubtful). 


NEW GUINEA: Irian Jaya, SW.: Zipelius s.n. (in L, as annotated by Blume). 


Ecology. Not known. 


New account of Horsfieldia 2 101 


NOTES 


1. Horsfieldia aruana with mature male flowers is only known from the lecto- 
type, i.e., three unnumbered Zipelius collections in Leiden, which have membra- 
nous leaves, and which bear annotations by both Blume and Zippel ‘‘Myristica Sp. 
arb. 40-50, N. Guinea’’. Also deposited in Leiden are three duplicates of which two 
are unannotated. They form part of the syntype material including the Zippel- 
collections mentioned by Warburg under his H. novo-guineensis; see further, notes 
2 and 3. 


2. Warburg (p. 273) had rejected the name Myristica aruana Bl. and replaced it 
with a new name, H. novo-guineensis ‘Warb., as he considered the type of M. aruana 
a mixture and dubious. Later he in turn conceived H. novo-guineensis as a very 
variable species (it being based on a number of specimens), not indicating a 
holotype, citing M. aruana Bl. p.p. and M. nesophila Mig. in its synonymy, 
Annales II (i.e, not the one in Annales I). H. novo-guineensis is presently referred 
to H. irana, a new species. 


The specimens cited by Warburg for H. novo-guineensis (now the syntype) are 
the following: New Guinea (West), Zipelius s.n. (C’, several sheets of two different 
species); Beccari 684 (C’, fr.), 116 (CO) — (East), Sepik, Hollrung 657 (fr.) — Aru 
Isls., Moseley s.n. (fr.) — Dammar Isl., Riedel s.n. (fr.). 


The Zipelius specimens are all in Leiden and represent two different species. 
Warburg (p. 273) had alluded to the difference. Part of it is now chosen to typify 


the presently accepted name H. aruana and to lectotypify H. novo-guinensis as 
explained in note 3. 


The remaining material, marked Zipelius (139d) in Leiden (K, iso) is now 
accommodated in H. irana, the newly described species. 


I have not examined the Beccari specimens. Sinclair referred 1/6 to H. spicata 
and 684 to H. polyantha. Of these two species I have presently quite different ideas. 


Hollrung 657 was referred by Sinclair to H. spicata as well but I consider it to be 
H. pilifera or H. laevigata although its fruits are intermediate in size, 16-18 mm 
long, and the twigs tend to be ridged. 


Sinclair had not mentioned Moseley s.n., from the Aru Isls., in K, and I have not 
seen it. 


Riedel s.n. from the Dammar Isls is H. smithii. 


3. The name M. aruana BI. was considered as dubious and rejected by Warburg 
(p. 273), being based on Rumph’s “‘Palala aruana”’ (Herb. Amboin., Auct. p. 56) 
whereas he regarded the Zipelius specimens too different from Rumphius’s de- 
scriptions. I agree with Sinclair (l.c.) who accepts Blume’s name typified by the 
Zipelius collections (see note 1) as annotated by Blume at the Leiden Herbarium. 
Sinclair regarded the name aruana, thus typified, as synonymous with H. spicata in 
the wider sense. It will become clear that Warburg’s new name “‘novo-guineensis”’ 
remains with “‘aruana” as lecto-typified by Sinclair (pp. 122 & 123). Of the Zipelius 
collections, Sinclair had cited them under the number of “(139d)” as being H. 
iriana, newly proposed by him. 


102 Gard. Bull. Sing. 38(1) (1985) 


4. Doubtful specimens are Buwalda 4969 (K,L) from the Aru Isls. and b.b. 24414 
(L) from the Tanimbar Isls. The male flowers of both are immature and they may 
well belong to the type collection. Even more dubious is b.b. 24414 as the stamen 
column of its androecium apppears to be open for nearly %4 or 4%, and irregular 
whitish blotches on its leaves are similar to those usually found in H. irya and H. 
smithii. The anther columns of Buwalda 4969 are cleft to a depth of only Mo. 
Sinclair identified this as H. pilifera (at Kew) and as H. spicata (at Leiden), and 
b.b. 24414 as H. irya. 


5. Distribution. Because of the doubts concerning Buwalda 4969 discussed above 
and the reference by Blume to “‘Palala aruana’’, it cannot be ascertained that the 
present species occurs on the Aru Islands. 


29. Horsfieldia subtilis (Mig.) Warb. Fig. 1B(29); 15 g, h. 


Myristica subtilis Miq., Ann. Mus. Bot. Lugd.-Bat. 2,1 (1865) 50 — Horsfieldia subtilis (Miq.) Warb., 
Mon. Myrist. (1897) 286, t. 23 fig. 1-4; Markgraf, Bot. Jahrb. 67, 2 (1935) 152 — Type: ‘Zipelius (78) 
(WU; tro, LS, 7 y:): 


For further synonyms see under the varieties. 


Tree 2-10(-15) m. Twigs terete, not ridged, 1-3(-8) mm diam., early glabrescent, 
tomentum grey-brown, of hairs c. 0.1 mm long or less; bark finely striate, when 
older not flaking, lenticels fine, usually present. Leaves in 2 rows, membranous or 
as in var. calcarea + chartaceous, elliptic to oblong-lanceolate, broadest usually at 
or above the middle, 6-25(-28) x 2-9(-9.5) cm, base attenuate, tip acute-acuminate; 
upper surface drying dull olivaceous to brown, with or without fine paler dots, 
lower surface very early glabrescent, hairs 0.1 mm or less; without larger dark- 
coloured dots, the nerves darker and contrasting in colour or not; midrib above 
flattish or slightly raised; nerves 6-16 pairs, above thin and flat or raised, beneath 
with the submarginal arches faint or distinct; tertiary veins forming a lax network, 
usually indistinct on both surfaces; petiole 5-13 xX 1.2.5 mm; leaf bud 6-12 x 1.5-2 
mm, with hairs c. 0.1 mm. Inflorescences in & 1-2(-3) times ramified, common 
peduncle up to 10 mm long, rather few- to many-flowered, 2-8(-9) < 1.5-6 cm, in 
Q: 2-5(-8) cm long; glabrescent or with sparse tomentum of scattered stellate-scaly 
hairs 0.1-0.2 mm long; bracts 0.5-2 mm long, glabrescent, with fimbriate margins, 
caducous. Flowers in loose clusters of 1-8(-10) each; perianths 2-valved, glabrous, 
pedicel slender, glabrous, at base not articulated. Male perianth in lateral view 
circular or more or less broadly transversely sub-elipsoid, or broadly ob-triangular, 
usually distinctly flattened, and collapsing on drying, 1.3-2.4 x 1.8-3 mm, upper part 
generally subtruncate or (broadly) rounded, at base subtruncate to short-cuneate, 
pedicel not tapering, slender, 1-3 mm long, perianth at anthesis split to c. %4-Ya(- 
Yr), valves 0.1-0.2(in var. acuta up to ¥%) mm thick. Androecium laterally 
flattened towards the top, at base broadened and usually almost cylindrical, lateral 
view subquadrangular in outline, i.e., broadly rounded or subtruncate above, the 
androecium nearly filling the perianth, c. (0.7-)1-1.5(-1.7) x 1.4-1.6 mm; anthers 
(9-) 10-12, faintly septate when young, + erect, free portions at apex up to 0.1 mm 
long, anther column at apex narrowly hollowed for c. ¥%4-’3; androphore + slender, 
distinct, 0.2-0.5 mm iong, sometimes + hidden by the sagged anthers. Female 
perianth broadly ellipsoid to ovoid, or subglobose, 1.8-2.5 xX 2-2.5 mm, split at 
anthesis to c. 14, valves c. 0.2.mm (at base of perianth 0.3-0.6 mm) thick, pedicel 
1-5 mm long; ovary avoid, 1.1-1.5 x 0.8-1.1 mm, glabrous, style with minutely 
bilobed stigma c. 0.2 mm long. Fruits (1-) 5-15 per infructescence, either globose or 
subglobose, 0.9-1.2(-1.3) * 0.8-1.1(-1.3) cm (pseudostalk up to 1 mm), or in some 


New account of Horsfieldia 2 103 


varieties with larger ones rather ellipsold, 1.4-1.9 x 1.1-1.4 cm, with the top 
rounded to acutish, base rounded, without or with pseudostalk up to 3 mm; 
glabrous, drying blackish, without or with minute paler tubercles or lenticels; dry 
valves c. 1(-2) mm thick; seed subglobose to ellipsoid; stalk 1-7 mm; perianth not 
persisting. 


Distrubution. Aru. Isls., the whole of New Guinea. 


In order to accommodate a number of specimens with the fruits distinctly larger 
than the majority, local varieties are recognized. 


KEY TO THE VARIETIES 


la. Male perianth 2-3 mm wide. Fruits globose or short-ellipsoid, 9-12(-13) mm long incl. pseudostalk 
ok I 66 ie RB Te 0 ps a Oe ae AOE doe ear oe Ae, Ue Oe ee Ps te a. var. subtilis 


b. Male perianth c. 2.5-3 mm wide (always?). Fruits short-ellipsoid, 14-19(-20) mm long incl. 
a OR Ee 0 at sci Asien oe does pe. iain Ses ain righlns + gs Ba us Avene nse oye H om AE oe nd p owe Oo a Sens 2 


2a. Leaves chartaceous, elliptic, 6-9 cm long. Pseudostalk of fruit 2-3 mm long. Limestone area, SW. 


ee nce Feomts £ Zeon meal eet 2. RIS. Fe wk b. var. calcarea 

b Leaves coriaceous or membranous, elliptic-oblong, 10-22 cm long. Pseudostalk of fruit 0-3 mm long. 
oo lee ASUS i Sa UE I i J RS PA Ra NA 3 

3a. Leaves membranous. Perianth (Q) glabrous inside ...................cecceceeceeceee eee eeeee eens c. var. aucta 
eee = COR aCeous. Feriantn,( OD). Wally MISIGE «° 320., van s+ cpineosnedeare onevescieeedericoeree. d. var. rostrata 
a. var. subtilis Fig. 1B(29); 15 g,h 


Horsfieldia subtilis var. subtilis: Sinclair, Gard. Bull. Sing. 28 (1975) 132. 

H. aruensis Warb., Mon. Myrist. (1897) 284, t. 23 fig. 1-3; Markgraf, Bot. Jahrb. 67, 2 (1935) 154 — 
Myristica aruensis (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 85 — Type: Beccari s.n. (FI Acc. 
Nos. 7622, 7622 A-C, 7623, n.v.). 

H. lauterbachii Warb., Mon. Myrist. (1897) 285, t. 23 fig. 1-2; Schumann & Lauterbach, Fl. Deutsch. 
Schutzgeb. (1900) 324; Pulle, Nova Guinea 8 (1912) 635; Markgraf, J. Arn. Arb. 10, 2 (1929) 213; 
Bot. Jahrb. 67, 2 (1935) 153 — Type: Lauterbach 805 (B, + BRSL, n.v.). 


H. ramuensis Warb. in K. Sch. & Laut., Fl. Deutsch. Schutzgeb., Nachtr. (1905) 266 — Type: Rodatz 
& Klink 20, 24 (both B 7). 


H. globularia auct. non (B\.) Warb.: K.Sch. & Laut., Fl. Deutsch. Schutzgeb. (1900) 324. 


H. nesophila auct. non (Miq.) Warb.: Pulle, Nova Guinea 8 (1912) 635. 


Leaves membranous, elliptic to oblong. Male perianths 1.8-3 mm wide. Fruits 
(sub)globose, c. 9-12(-13) x 8-11 mm, incl. the pseudostalk 0-1 mm long. 


Distribution. As for the species. 
ARU ISLS. Buwalda 4970; Jensen 255. 


NEW GUINEA. Irian Jaya: Aet 2, 313, 382: Aet & Idjan 554; Astarip 63, 715; BW 2362, 2464, 3517, 
3518, 4340, 4761, 4834, 4837, 4942, 6056, 6235, 6796, 8377, 10278, 10628, 11388, 12255, 13524, 13576; 
Docters van Leeuwen 9122, 9611, 9698, 9767, 10702, 11066, 11067, 11222; Gjellerup 11, 273; ljiri & 
Niimura 53; Kostermans 2668, 2670, 2686, 2802, 2903, 4744; Lam 770; Pymans 7383; Pleyte 1006; Pulle 


104 Gard. Bull. Sing. 38(1) (1985) 


52, 1239; von Rémer 676; van Royen 3473, 3560, 4017, 4745, (& Sleumer) 6694; Soegeng 366; Teysmann 
7566; Versteeg 1140, 1568, 1612, 1616, 1814; Zipelius 78 — Papua New Guinea: Brass 1414, 28938; Carr 
11545, 11629, 12575, 12820, 16281; Craven & Schodde 840, 1012; Darbyshire 1024; Hartley T.G.H. 
10710, 11328; Hoogland (& Womersley) 3245, 3503, 3617, 4208, 4583, (& Craven) 10475; Jacobs 9129, 
9252; Kanis 1031; LAE 51573, 61106, 61231, 63002, 66272, 70255, 70256, 70455, 73949, 76136; NGF 
3890, 4553, 8232, 10378, 13066, 13146, 16068, 16075, 17775, 18308, 18433, 18436, 23575, 24844, 31731, 
32822, 33410, 35340, 35427, 35449, 35472, 41108, 41878, 43692, 43693; Pullen 1074, 7280, 7419, 8139; 
Schodde 2590, 2910, 2968, 3066; Womersley 3890, (& Simmonds) 5080. 


Ecology. Understorey tree of primary and secondary forests; dry and marshy 
forest, but often tidal (fresh water) or riverine; on alluvia, clayey soil, sandy clay, 
also on limestone or coral soils; 0-800 m. Flowers and fruits throughout the year. 
Stems once reported as inhabited by ants. 


Vernacular names. Aitobi (Aru Isls.), Airawikoepata (West N.G., Tisa), Ben- 
doei (Vogelkop, Hattam lang.), Boskomok (East N.G., Western Prov., Oriomo 
R.), linapo (Uruaruh lang., Gulf Prov.), Mag (E. N.G. Gulf Prov., Daru lang.), 
Mangaifa (Papua, Centr. Prov.), Njet (W. N.G., Kebar lang.), Oara (Papua, 
South Vieya), Peh (begie) (Div. South N.G., Digoel R., Awjoe lang.), Reng- 
kéferék (W. N.G., Beriat, Tahid land.), Rewwoh f(W. N.G., Fak-Fak, Argoeni 
lang.), Roman (E. N.G., Pt. Moresby, Centr. Prov., Waria lang.), Suri (E. N.G., 
Sepik Prov., Waskuk lang.), Tabwi (E. N.G., Sepik Prov., Wagu lang.), Torua (E. 
N.G./Papua, North. Prov., Baruga lang.). 


Uses. Once recorded that the leaves and twigs were burnt as a mosquito repel- 
lent. 


NOTES 


1. Fieldnotes. Low understorey tree, usually 3-5 m tall. Bole straight; bark 
greyish black or grey-brown, finely longitudinally fissured, with broadened len- 
ticels. Branches often horizontal or drooping. Exudate pinkish, or colourless and 
turning reddish. Wood straw-coloured, usually mottled with reddish streaks. 
Perianth yellow, rarely orange-yellow. Fruits greenish-yellow, yellow, or (yellow) 
orange, aril orange or red. 


2. Deviating specimens. Van Royen & Sleumer 6694 from a coral cliff near 
Mankokwari (Vogelkop) somewhat deviates in the relatively narrow perianth, + 
longer than broad, c. 2.4 xX 2.2 mm. Versteeg 1612 from SW. New Guinea has 
relatively large fruits, c. 1.3 x 1.1-1.3 cm. 


3. The type-variety as presently accepted largely agrees with Sinclair’s H. subtilis 
var. subtilis. Of the other two varieties accepted by him, are var. rostrata (Mgkf.) 
and var. schlechteri (Warb.), the latter presently treated as a different species, 
distinguished by characters different from those used in his key to the varieties. 


b. var. calcarea de Wilde, var. nov. 


Differt a H. subtilis var. subtilis perianthiis masculis latioribus atque fructibus maioribus, c. 1.5-2 cm 
longis, stipitibus 2-3 mm longis, foliis chartaceis, usque ad 10 cm longis. — Type: Vink BW 15270 (L; 
iso: K; A, BO, BRI, CANB, LAE & US, n.v.). 


Leaves thinly chartaceous, elliptic, 6-9 x 2.5-4 cm, at the apex proportionally 
long-acute-acuminate for 1-1.5 cm. Male flowers not seen. Fruits ellipsoid, 17-19 x 


New account of Horsfieldia 2 105 


12-14 mm, incl. pseudostalk 2-3 mm long. 


Distribution. West New Guinea, SW. Vogelkop Penins. 


IRIAN JAYA. Vogelkop Penins.: (Versteegh) BW. 7432; (Vink) BW. 15270. 


Ecology. Secondary forest on limestone rock with thin clay cover; 220-300 m alt. 
Fruits in March and May. Female flowers in May. 


Vernacular name. Baiwach, Hafringee (Maibrat lang.). 
NOTES 
1. Recorded as a shrub, 5 m tall; rather common. Ripe fruits orange. 


2. Judging from the aspect after drying, the pericarps of the fruits in Vink BW 
15270 suggested they were rather fleshy in the fresh state and drying left the 
pseudostalks distinct, c. 3 mm long. 


c. var. aucta de Wilde, var. nov. 


A Horsfieldia subtilis typica differt fructibus maioribus, a H. subtilis var. calcarea foliis maioribus 
membranaceis — Type: Jacobs 8972 (L; iso: K). 


Leaves membranous, elliptic-oblong, 11-20 x 3-7.5 cm, apex for 1-1.5 cm acute- 
acuminate. Male perianth obtriangular, c. 2.5-3 mm wide (always?). Fruits short- 
ellipsoid, 15-19(-20) x 11-15 mm including the up to 2.5-mm long pseudostalk. 


Distribution. Papua New Guinea; possibly also near Manokwari, Vogelkop 
Penins., W. New Guinea (see notes). 


NEW GUINEA. Irian Jaya, Vogelkop: (Koster) BW 4340 (doubtful). Papua New Guinea: (Western 
Dist.) NGF. 42815; (S. Highlands) Jacobs 8711, 8972, 9053, 9071, 9071-A; (Central Distr.) Kanis 1328. 


Ecology. Montane primary and secondary rain forest, on well-drained volcanic 
soil, or peaty soil; at 600-1000 m. Flowers in September and October, fruits from 
July to October. 


NOTES 


1. Shrub or low tree, 3-8 m. Male flowers fleshy, dark yellow. Fruits glossy 
orange, hard; aril dark orange, or red, at the base black. 


2 The male flowers, known from Jacobs 9071, are stouter and have a thicker 
perianth as compared with those of var. subtilis. In Jacobs 9071 the perianth is 
obtriangular, c. 3 x 3 mm, and split into 2 (or some into 3) valves only for the apical 
Ys-/4; lower down, the perianth wall is thick-fleshy, c. 0.5 mm thick. On Jacobs 
9071-A, a collection from a nearby tree with fruits measuring c. 16 x 14 mm when 
dry, Jacobs had remarked that in the fresh state the ‘‘seed (is) half the diameter of 
the fruit’. The dry seed measures about 14 mm, and probably was not or only 
slightly larger in the fresh state; this means that fresh fruits were about 3 cm diam., 
i.e., the dry ones in the herbarium have shrunk to half. 


106 Gard. Bull. Sing. 38(1) (1985) 


_ The inflorescences in Jacobs 8711, with submature fruits, are 8-9 cm long; these 
are stouter than generally found in the type variety. 


3. Kanis 1328, from Moresby area, deviates by a more slender habit. Koster BW 
4340, from near Manokwari, Vogelkop (West New Guinea) at an altitude of 
c. 150 m. probably does not belong here. Moreover, its fruits are c. 14 X 12 mm, 
considerably larger than all the rest seen of var. subtilis from the same area. 


d. var. rostrata (Mkegf.) Sinclair 


Horsfieldia subtilis var. rostrata (Mkgf.) Sinclair, Gard. Bull. Sing. 28 (1975) 136 — Horsfieldia rostrata 
Mkgf, Bot. Jahrb. 67, 2 (1935) 152 — Type: Ledermann 8916 (B, +; iso: SING, n.v.). 


Leaves chartaceous to coriaceous, cuneate-obovate, c. 9-13 x 2-4.5 cm, tip 
shortly acute-acuminate. Male flowers not known. Female perianth pilose inside 
(see notes). Fruits ellipsoid, top rostrate, c. 20.x 12 mm, including the 2-4-mm long 
pseudostalk. 


Distribution. NE. Papua New Guinea: Sepik Prov., Etappenberg; known only 
from the type. 


Ecology. Mossy montane forest with much Agathis, c. 850 alt. Female flowers 
and fruits in October. 


NOTES 


The holotype of H. rostrata, containing 9 flowers and mature fruit, was lost in B; 
an isotype is in SING and was examined by Sinclair. He maintained H. rostrata as 
a variety under H. subtilis. According to him, the SING specimen contains a single 
infructescence with two fruits, each measuring c. 1.4-1.5 x 1 cm; the fruits have ac. 
2-mm beak, and pseudo-stalks 3-4 mm long. According to this description the 
foregoing new var. aucta appears almost entirely identical with var. rostrata. 
However, Markgraf, I.c. p. 153, describes the perianth of the female flowers as 
pilose inside; the ovary and fruit as glabrous. The hairiness of the inner side of the 
perianth would render the Ledermann 8916 collection highly remarkable, and 
consequently I have provisionally kept it as a separate taxon. The inner hairy 
surface of the perianth seems reminiscent of Endocomia macrocoma, but this has 
three perianth lobes. 


30. Horsfieldia schlechteri Warb. Fig. 1B(30); 15 a-f. 


Horsfieldia schlechteri Warb. in Schum. & Lauterbach, Nachtr. z. Fl. Deutsch. Schutzgeb. Siidsee 
(1905) 267; Markgraf, Bot. Jahrb. 67, 2 (1935) 153 — H. subtilis var. schlechteri (Warb.) Sinclair, 
Gard. Bull. Sing. 28 (1975) 137 — Type: (Torricelli Mts.) Schlechter 14500 (B, + iso: BM, K, P; BO, 
WRCL, G, n.v.). 


Tree or shrub, 3-15 m. Twigs terete, not ridged, towards apex 1-4(-6) mm diam.., 
early glabrescent, tomentum rusty-grey, with hairs c. 0.1 mm long; bark finely 
striate, when older not flaking; without or with few lenticels. Leaves in 2 rows, 
membranous or thinly chartaceous, elliptic-oblong to oblong-lanceolate, + paral- 
lel-sided or note, 6-19 x 1.06-6.5 cm, base attenuate, tip acute-accuminate; upper 
surface drying olivaceous to dark brown, finely paler pustulate or not, lower 
surface early glabrescent, the hairs 0.1 mm or less, without larger brown dots, the 


Fig. 15. 


Jb 


Horsfieldia schlechteri Warb. a, leafy twig with male inflorescences, X ‘2; b, mature male 
flower, X 12; c, ditto, opened, showing stiped androecium, x 12; d, opened mature female 
flower, showing glabrous ovary, X 12; e, portion of twig with infructescence with mature 
fruits, X 2; f, mature fruit of a different specimen, xX 2. — Horsfieldia subtilis Miq.) Warb. 
var subtilis: g, opened male flower, showing androecium, androphore hidden by the anthers, 
x 12; A, twig portion with mature infructescence. — a-c, from Kostermans & Soegeng 359; d 
& e, from BW 4307; f, from BW 2900; g, from LAE 70256; h from Hoogland 3503. 


107 


108 Gard. Bull. Sing. 38(1) (1985) 


nerves little contrasting; midrib slender, raised from both surfaces; nerves 6-14 
pairs, raised or flattish above, indistinct; tertiary veins forming a lax network very 
indistinct on both surfaces; petiole 5-12 x 0.7-1.5 mm; leaf bud c. 8 X 1 mm, with 
hairs c. 0.1 mm long. Inflorescences glabrescent or with rather sparse to dense 
tomentum of stellate hairs 0.1 mm long or less, sometimes extending to the 
pedicels; in C’: (1-)2(or 3) times ramified, 1.5-7 x 1-4 cm, common peduncle 3-15 
mm, slender; in Q: few-flowered, 1-4 cm long; bracts 0.5-1 mm long, densely 
woolly pubescent, caducous. Flowers solitary or 2 or 3 together, perianths 2-valved, 
glabrous; pedicels slender, at base inarticulate. Male perianth (broadly obovoid 
or) subglobose, (1-) 1.3-1.5 x (1-) 1.5-2 mm, not or only little laterally compressed, 
when dry wrinkled but not or slightly collapsed, rounded above, shortly tapered at 
base; pedicel slender, (1-)2-3.5 mm long, slightly broadened towards the perianth, 
glabrous or with some scattered minute hairs at base; perianth at anthesis split to c. 
Y4(-'2), valves 0.1-0.2 mm thick. Androecium + flattened especially in the upper 
part, thickish at base, broadly ellipsoid in outline, above and at base broadly 
rounded, c. 0.6-0.8 xX 0.9-1.0 mm; anthers 8, mutually appressed, free apices 0-0.1 
mm; androphore 0).4-0.5 mm; anther column at apex narrowly excavated for c. %. 
Female perianth ellipsoid-obovoid, c. 1.5-2 xX 1.5 mm, split at anthesis to c. %, 
valves 0.1(-0.2) mm thick, pedicel 1.5-2 mm long; ovary ellipsoid, c. 1-1.2 x 0.8 
mm, glabrous or with a few minute hairs at apex (see notes), stigma minutely 2- 
lobulate, c. 0.1 mm long. Fruits 1(-2) per infructescence, ellipsoid, top rounded, 
not pointed or for only c. 1 mm, base rounded to short-attenuate into a pseudostalk 
(1.5-) 2-6 mm long, excl. pseudostalk 1.3-2.0(-2.5) x 0.9-1.2 cm, glabrous, drying 
blackish, with or without paler pustules (or lenticels), dry valves 1-2 mm thick; stalk 
5-10 mm long; perianth not persisting. 


Distribution. New Guinea: Irian Jaya, Jayapura Dist.; N. Papua New Guinea, 
W. Sepik Prov. 


NEW GUINEA. Irian Jaya: bb 25083; BW 2900, 3681, 4064, 4307, 5522; Kostermans & Soegeng 235, 
359; van Royen & Sleumer 6219, 6455 — Papua New Guinea: Hoogland & Craven 10703; NGF 13293, 
18953, 39223, 48230, 48297; Pullen 1514; Schlechter 14500. 


Ecology. Forest on stony slopes, foothills, ridges; mossy forest, lowland and 
lower montane forest; on stony clay, sandy soil; 20-500 m alt. Flowers and fruits 
throughout the year. 


Vernacular names. Cheem (Sepik Prov.), Guma (Waskuk lang., Sepik Prov.), 
Medal (Wagu lang., Sepik Prov.). 


NOTES 


1. Fieldnotes. Bark dark brown or blackish, with longitudinal fissures. Perianths 
yellow or orange-yellow. Fruits green-yellow, yellow, or (yellow-)orange; aril red. 


2. BW4307 (Hollandia, 400 m) has markedly large fruits, c. 2.5 x 1.2 cm incl. the 
5-6-mm long pseudostalk. The specimen Kalkman 3681, from the same area, at 50 
m, has fruits c. 1.3 x 0.9 cm, with only a short pseudostalk (1-) 1.5 mm; its ovaries 
and very young fruits have a few minute hairs towards the apex; it approaches 
certain specimens of H. subtilis. 


New account of Horsfieldia 2 109 


3. H. schlechteri, as accepted presently, largely agrees with Sinclair’s H. subtilis 
var. schlechteri. However, specimens cited by him from outside the Jayapura/Sepik 
area are presently referred to H. subtilis. 


31. Horsfieldia basifissa de Wilde, sp. nov. Fig. 1B(31) 
Horsfieldia polyantha auct. non Warb.: Sinclair, Gard. Bull. Sing. 28 (1975) 95, p.p. 


A Horsfielda speciebus quoad perianthia 2-valvata sese similibus, differt ramulis non-cristatis, 
perianthiis masculis sub anthesi usque ad basin divisa, androecio excavato per 3-2, atque fructibus 
glabris (sub)globosis. — Type: White NGF 10242 (L). 


Tree 10-25 m. Twigs terete, faintly ridged or not, towards the apex 2-4(-8) mm 
diam., early glabrescent, tomentum grey-brown, with hairs c. 0.1 mm; bark finely 
striate, when older not flaking; lenticels inconspicuous. Leaves in 2 rows, membra- 
nous to thinly chartaceous, elliptic-oblong to oblong-lanceolate, broadest usually 
above the middle, 10-22 x 3-8 cm, base attenuate, top acute-acuminate; upper 
surface drying olivaceous to brown (often with paler markings, possibly caused by 
calcium agglomerations), not or faintly, minutely, more palely pustulate, lower 
surface glabrescent, hairs c. 0.1 mm; without larger dark brown dots; midrib above 
flattish; nerves 10-15 pairs not particularly contrasting, above thin and flattish or 
sunken, beneath with the marginal arches not very regular nor prominent; tertiary 
veins forming a lax network, rather faint; petioles 5-10 x 1.5-2.5 mm; leaf bud c. 10 
x 1.5 mm, with hairs c. 0.1 mm. Inflorescences in 0:3(-4) times ramified, many- 
flowered, 4-10 xX 2.5-6 cm, common peduncle 2-20 mm long; in 2: c. 5 X 3.5 cm; 
densely to sparsely pubescent with stellate hairs 0.1-0.2 mm; bracts elliptic-oblong, 
acute, 1-2(-4) mm long, caducous. Flowers generally 1-3 together; perianth 2- 
valved, glabrescent except at the very base, pedicel thinly pubescent with hairs c. 
0.1 mm long, at base not articulated. Male perianth as seen laterally subcircular, 
slightly broader than long, only little laterally compressed, not or but slightly 
collapsed on drying, 2.2-2.7 x 2.6-3 mm, upper and basal part broadly rounded, 
pedicel not tapering, 1.5-3 mm long; perianth at anthesis cleft to the base, valves 
0.1-0.2 mm thick. Androecium laterally much flattened, above broadly rounded, c. 
1.5-1.7 x 2.0 mm; anthers 12-14(-16), distinctly septate when immature, erect, free 
portions at apex c. 0.1 mm long, anther column at apex narrowly hollowed for 
Y3-*/3; androphore 0-0.1 mm, broadly attached. Female perianth (immature flowers 
seen only) broadly ovoid, c. 1.5 X 1.4 mm, split at anthesis nearly to the base, 
valves c. 0.2-0.3 mm thick, pedicel 1.5-2 mm long; ovary ovoid, c. 1.1 xX 0.6 mm, 
glabrous, style and stigma small, minutely 2-lobed. Fruits 1-20 per infructescence, 
globose or subglobose, 1.1-1.4 cm diam., glabrous, drying light to dark brown, with 
or without coarse, paler-coloured lenticels or warts; dry valves c. 1.5-3 mm thick, of 
woody-granular structure; seed ellipsoid; stalk 3-4 mm; perianth not persisting. 


Distribution. New Guinea: NE. Irian Jaya; N. Papua New Guinea (Sepik, 
Madang Prov.). 


NEW GUINEA. Irian Jaya: (Schram) BW 2665; (Kalkman) BW 3455 — Papua New Guinea: 
Hoogland & Craven 10215; NGF 10242, 26953, 32824, Pullen 1896; Saunders 198, 958; Schlechter 18302; 
Womersley 3798, 3821, 3899. 


Ecology. Primary and secondary forest, marshy forest, locally common; also 
recorded from Pometia-Intsia forest on clays and marls; 0-200 m alt. Flowers in 
September, fruits in March, June, and October. 


110 Gard. Bull. Sing. 38(1) (1985) 


Vernacular names. Euoe (Sko lang., Hollandia); Ilis (Jal, Madang Prov.); Num- 
ba (Angorami, Sepik Prov.). 


NOTES 


1. Fieldnotes. Slender tree, branches horizontal. Flowers yellow; fruit green, 
turning orange. 


2. Apart from H. angulata (see the notes under that species) it is possibly closely 
related to H. parviflora, because of the glabrous fruits. The fruits are globose, and 
often are very similar to those of H. pilifera or H. sinclairii; in these two species, 
however, the fruits are always hairy, at least towards the base. See also note 4. 


3 .Many of the specimens accepted in the present new species were identified by 
Sinclair (p. 97) as H. polyantha, with H. novoguineenis as a synonym. The speci- 
mens of the syntype of H. polyantha, Beccari 7619, 7619 A, not seen by me, 
however, most likely belong to Knema laevigata. 


The syntypes of H. novoguineensis Warb. are very heterogeneous; the two 
lectosyntypes (Sinclair, p. 95), Beccari 684 (not seen), and Zipelius 139-d belong to 
H. iriana; a third syntype, Hollrung 657 (fruits) belongs to H. pilifera (a species 
close to H. laevigata) whereas other syntypes belong to yet other species. 


4. The new species has much in common with H. laevigata var. novobritannica, 
which has the androecium also deeply hollowed inside; the latter has, however, a 
more hairy perianth. Of var. novobritannica, the female flowers are not known, 
but it has globose fruits larger than those of H. basifissa, and they are somewhat 
hairy at the base. The present new species is characterized by the subglabrous male 
flowers with a very deeply cleft perianth, glabrous ovary and glabrous globose 
fruits; it could be confused with H. pilifera and H. sinclairii which also may have 
globose fruits, but are never glabrous. 


32. Horsfieldia sinclairii de Wilde, sp. nov., Fig. 1B(32) 


Horsfieldia erubescens Sincl., in sched. (Gard. Bull. Sing. 28, 1975, 6-7) — H. australiana auct. non S.T. 
Blake: Sinclair, Gard. Bull. Sing. 28 (1975) 6, p.p. — Type in sched.: Womersley & Brass. NGF 8664 
(SING, n.v; iso: BM, K, L; A, BO, BRI, CANB, LAE & NSW, n.v. 


Perianthia mascula parva, subglobosa, c. 1.5-2 mm diam., glabra, sub anthesi usque ad 1 divisa, 
antheribus 6-10, androphoro usque ad c. ¥3 excavato, ovario pubescente, fructibus globosis vel breviter 
ellipsoideis, 1.5-2.5 cm longis, in siccitate nigrescentibus, pericarpio sicco 4-6 mm crasso. — Type: 
(Streimann & Katik NGF 28886 (L, iso: K). 


Tree, 4-25 m. Twigs terete, not ridged, towards apex 1.5-3(-6) mm diam., early 
glabrescent, tomentum brown, of hairs c. 0.1 mm; bark finely striate, when older 
not flaking; lenticels mostly inconspicuous. Leaves in 2 rows, membranous, elliptic- 
oblong to oblong-lanceolate, broadest at or above the middle, 6-14 x 1.7-4.5 cm, 
base attenuate, tip acute-acuminate, upper surface drying light to dark brown, 
sometimes with paler markings, minutely, more palely pustulate or not; lower 
surface glabrescent, hairs c. 0.1 mm, without larger dark brown dots, the nerves 
usually not much contrasting; midrib slender, flat above, often reddish tinged and 
contrasting below; nerves 6-14 pairs, thin and flat above, beneath inconspicuous, 
marginal nerve faint; tertiary veins forming a lax network, inconspicuous; petiole 
6-15 x 0.8-1.5 mm, leaf bud 8-15 x 1-2 mm with hairs c. 0.1 mm long. Inflor- 


wee . 


New account of Horsfieldia 2 111 


escences sparsely minutely hairy to subglabrous, hairs c. 0.1 mm; in CG: many- 
flowered, 2-4 times ramified, 2.5-8 < 1.5-6 cm, common peduncle 2-10 mm long; in 
Q: up to 5(-10) X 4.cm; bracts 0.5-2.5 mm long, caducous. Flowers in loose clusters 
2-5 each; perianths 2-valved (in some specimens a rather high percentage of 
3-valved perianths), glabrous; pedicels slender, glabrous, at base inarticulate. Male 
perianth subcircular (rarely slightly broader than long), slightly laterally compress- 
ed, not or somewhat collapsed on drying, 1.1-2.0 x 1.5-1.8 mm, broadly rounded 
above, rounded to broadly rounded below, pedicel not tapering, slender, 0.6-1.5 
mm long; perianth at anthesis cleft to c. ¥2-way, valves 0.1-0.2 mm thick. 
Androecium + flattened, in lateral view broadly circular to subobtriangular, above 
broadly rounded, c. 0.6-1.4 x 1-1.3 mm; anthers 6-10, septate when immature, free 
portions at apex rather conspicuous, 0.1-0.3 mm long, the anther-bearing 
column at apex not or only moderately hollowed up to c. 3; androphore narrow, 
0-0.3 mm long. Female perianth much larger than in OC, c. 2-2.4 x 1.8-2.2 mm, 
ellipsoid-ovoid, at anthesis split to c. 43, valves c. 0.2-0.3 mm thick, pedicel c. 1 mm 
long; ovary globose, c. 1.6 mm diam., densely minutely pubescent, style + absent, 
stigma distinctly 2-lobed, c. 0.2 mm long. Fruits 1-5(-10?) per infructescence, 
globose to short-ellipsoid, or obovoid, top rounded, base rounded or contracted 
into a pseudostalk up to 2mm, rather distinctly ridged or not, drying dark brown to 
blackish, 1.5-2.5 x 1.5-2.0 cm, glabrescent but often with remnants of tomentum 
towards base (lens!), with or without only a few coarse lenticels or tubercles; dry 
valves 4-6 mm thick, woody; stalk 1-4 mm long; perianth not persisting. 


Distribution. Papua New Guinea: Madang Prov., Morobe Prov., Northern 


Prov., Milne Bay Prov. (incl. Fergussion Isl. and Normanby Isl.), Central Prov.. 
Gulf Prov. 


PAPUA NEW GUINEA: Brass 21821, 21912, 21996, 21997, 25503; Carr 12398; Clemens 877; Hartley 
T.G.H. 9968, 10421; Hoogland 5139, 8957 (deviating); Hoogland & Mc Donald 3516; Kanis 1107; LAE 
60239, 67162, 68817, 70202, 70272, 72475; NGF 8247, 8664, 28886; Saunders 528; Schodde 5638, (& 
Craven) 4366 (deviating). 


Ecology. Primary and disturbed lowland and mountainous rain forest, flood- 
plain forest, on slopes, ridges, also along creeks on stony places. Understorey tree; 
found on Castanopsis-dominated ridge, and in Anisoptera-Hopea-dominated 
forest. Altitude 0-950 m. Flowers mainly in March, April & June, fruits mainly in 
July and October. 


Vernacular names. Gaigihab (Dumpu; Madang Prov.), Hamana (Orokaiva 
lang., Mumini; Northern Prov.) Posiposi (Milne Bay Prov.), Saksak (Amele; 
Madang Prov.). 


NOTES 


1. Fieldnotes. Flowers creamy, yellow, or yellow-orange; twice recorded as tra- 
grant. Fruits glossy green, turning yellow to orange. Once recorded as with buttres- 
ses 1 x 1 ft. Bark rough, fissured or peeling off in irregular flakes leaving concave 
depressions. Wood cream- or straw-coloured or brown. Bark with reddish exudate. 


2. Relationship. Characterized by the slender twigs, smallish thin leaves with 
often reddish-tinged midrib beneath, small subglobose glabrous male perianths, 
the much larger female flowers with pubescent ovary, and the short-ellipsoid fruits 
drying dark brown or blackish, usually with 4-6-mm thick woody pericarp. The 


£2 Gard. Bull. Sing. 38(1) (1985) 


fruits may be confused with those of H. laevigata, especially with those of certain 
specimens from SW. New Guinea, which have rather similar pericarps but much 
larger leaves. In the present species the fruits have often become quite glabrous, 
but younger fruits (and ovaries) are pubescent; often minute hairs can be seen close 
to the bases of old fruits. 


3. The present new species was at first recognized by Sinclair, who named the 
sheets and in his manuscript as H. erubescens. A type (NGF 8664) was also 
designated. Shortly before his death Sinclair must have been of the opinion that his 
new species was conspecific with H. australiana, hence the use of H. australiana in 
the posthumous edition of his manuscript. 


4. In general habit H. australiana resembles but differs in various ways, especial- 
ly in the androecium. 


5. The deviating specimens Hoogland 8957 and Schodde & Craven 4366, both 
with male flowers, key out in the vicinity of H. sinclairii, but apparently are not 
conspecific. They rather agree with H. sinclairii in general appearance, but differ in 
their larger male flowers and in the pedicel being thinly minutely pubescent, also 
the basal part of the perianth in Schodde & Craven 4366. The latter collection is in 
these respects and in its rather elongate perianths reminiscent of H. pilifera, but 
for the larger flowers. Hoogland 8957, from Morobe Prov., c. 900 m, has male 
perianths subcircular in lateral view, c. 2.4 mm diam., cleft at anthesis to c. 3; 
there are c. 10 anthers, the pedicel is minutely sparingly pubescent. Schodde & 
Craven 4366, from Gulf Prov. at c. 300 m, has the male perianths rather elongate, 
c. 2.5(-3.0) x 1.8-2.0 mm, cleft at anthesis to c. 12-7, anthers c. 6 or 7, pedicel and 
basal part of perianth minutely pubescent. Probably these specimens represent 
separate taxa. 


33. Horsfieldia psilantha de Wilde, sp. nov. Fig. 1B(33) 


Perianthium masculum lateraliter compressum, aspectu laterali subcirculare, 2.5-3.5 mm diam., 
glabrum, pedicello gracili, haud attenuato. Infructescentiae usque as 15 cm longae, laxae, fructibus 
ellipsoideis, 17-22 mm longis, minute pubescentibus — Type: Long Island,  fl., Womersley NGF 43642 
Gb, iso: 3); 


Tree 5-25 m. Twigs terete, not ridged, towards the apex 3-6(-15, in fruiting twigs) 
mm diam., rather early glabrescent, tomentum reddish or grey-brown, composed 
of hairs c. 0.1-0.3 mm; bark finely striate, lenticellate, when older not flaking. 
Leaves in 2 rows, membranous, oblong-lanceolate to lanceolate, sometimes almost 
parallel-sided, 20-40 x 4.5-12.5 cm, base attenuate, tip acute-acuminate; upper 
surface drying olivaceous to green-brown, usually minutely pale-punctate; lower 
surface glabrescent or if leaves are younger, with scattered stellate hairs 0.1-0.3 mm 
on and near the midrib, without large dark dots, the nerves + greenish or reddish 
brown; midrib above flattish; nerves 14-24 pairs, above thin and flattish, beneath 
with the submarginal arches rather distinct, not very regularly shaped; tertiary 
veins forming a lax network, indistinct; petiole 5-20 x 2-3.5 mm; leaf bud 20-25 x 
2.5-3.5 mm, pubescent with hairs 0.1-0.3 mm. Inflorescences with rather thin 
woolly tomentum or stellate-dendroid hairs 0.2-0.4 mm; in CO and Q (when 
fruiting): 3 or 4 times ramified, many-flowered, 10-16 x 8-12 cm, common pedun- 
cle 10-40 mm; bracts not seer, caducous. Flowers in loose clusters of 2-5 each, 
perianths 2-valved, glabrous; pedicels slender, glabrous, at base inarticulate. 


Male perianth subcircular, somewhat laterally compressed, c.(2.0-) 2.5-3 x (2/5-) 


New account of Horsfieldia 2 113 


3-3.5(-4) mm, upper part broadly rounded, basal part rounded to short-attenuate, 
pedicel not tapering, (2-)3-4(-4.5) mm long; perianth at anthesis split to c. 12-4; 
valves c. 0.2-0.3 mm thick. Androecium much laterally flattened, rounded-truncate 
above. 1.4-1.8 X (1.5-)1.8-2.0(-2.2) mm; anthers c. 12-14, mutually appressed, not 
septate, erect, apices free for c. 0.1 mm; androphore up to 0.2 mm; anther column 
at apex narrowly hollowed for c. ¥4-2. Female flowers not seen. Fruits up to 10 per 
infructescence, ellipsoid, top and base obtuse to rounded, 1.7-2.2 x 1.2-1.7 cm, 
pubescent though sometimes hairs only remaining at the very base; hairs rusty, c. 
0.2 mm long; pericarp drying brown, without or with scattered small or fine 
lenticels or wartlets, 1-2 mm thick; stalk 2-8 mm long; perianth not persisting. 


Distribution. NE. New Guinea. Madang Prov.: Bagabag Isl., Long Isl; New 
Britian; New Ireland. 


PAPUA NEW GUINEA. New Britain (W. & E.): LAE 52132; NGF 21797, 21902, 30448, 41421 — 
New Ireland: Sands et al. 2047 — Madang Prov., S. Bagabag Isl.: NGF 42229A — Long Isl: LAE 55033; 
NGF 42361, 42390, 42398, 43640, 43642. 


Ecology. Forest (incl. beach), in shaded secondary forest; 0-200 m alt. Flowers in 
May and October, fruits throughout the year. 


NOTES 


1. Fieldnotes. Slender tree, branches often drooping, without or with a few 
buttress-roots. Bark blackish or dark grey-brown, longitudinally fissured; inner 
bark cream or pink, exudate pink or colourless; sap wood straw- or cream-coloured. 
Flowers orange-yellow. Fruit yellow to orange, aril orange. 


2. Related species are H. tuberculata, H. laevigata and H. whitmoret. H. tubercu- 
lata, variable and wide-spread, has similarly glabrous flowers, but the shape of the 
perianth is more tapered at the base, while in the present species it is more circular 
in lateral view and at the base not or but slightly tapered: H. tuberculata furth- 
ermore has glabrous fruits and ovaries. H. laevigata, a variable and wide-spread 
species as well, always has pubescent perianths, though sometimes only scattered 
hairs are present; it usually has smaller leaves, and the fruits usually have many 
more and coarser lenticel-like tubercles. The leaves of H. whitmorei, from the 
Solomon Isls, sometimes have similarly, rather regularly looping, marginal nerves, 
and similar fruits, but the male perianth is smaller, only c. 2 mm diam. or less, the 
base pubescent, also the pedicels, and the perianth cleft at anthesis to c. “10. NGF 
41421, particularly, from West New Britain, resembles H. whitmorei in general 
habit, especially in the marginal nerve and in leaf colour. 


3. Good representative specimens in fruit are LAE 55033 (from Long Isl) and 
NGF 42229A (Bagabag Isl). 


4. Some of the specimens have similarly large, branched and spreading in- 
fructescences as are the male inflorescences (known only from the type). Female 
flowering specimens are not known. 


34. Horsfieldia whitmorei Sinclair Fig. 1B(34); 16 


Horsfieldia whitmorei Sinclair, Gard. Bull. Sing. 27, 1 (1974) 135 — Type: Whitmore BSIP 1848 (SING; 
iso: K,L: LAE, n.v.). 


114 Gard. Bull. Sing. 38(1) (1985) 


Horsfieldia ‘palewensis’ (sphalm. palauensis) auct. non Kanehira: Whitmore, Guide For. Brit. Solom. 
Isl. (1966) 131, 186 in checklist. 


Tree 8-25 m. Twigs terete, faintly ridged or not, 2-5(-7) mm diam., + early 
glabrescent, tomentum often reddish-brown, composed of hairs 0.1-0.3(-0.4) mm; 
bark striate, when older not flaking; lenticels distinct or not. Leaves in 2 rows, 
membranous or rarely chartaceous, oblong to lanceolate, often almost parallel- 
sided, 9-30(-40) x 2-7(-9) cm, base short- to long-attenuate, tip acute-acuminate, 
upper surface drying dull brown to green brown, often minutely paler pustulate; 
lower surface largely glabrescent but often a few hairs remaining, hairs 0.1-0.4 mm, 
without larger brown dots, the nerves generally reddish-brown; midrib above 
flattish; nerves 18-26 pairs, above thin and sunken, beneath with the marginal 
arches very regular and distinct; tertiary veins forming a lax or fine network, 
distinct or not on both surfaces; petiole 10-15 x 1.5-3 mm; leaf bud 10-15 k 2 mm, 
with hairs 0.1-0.4 mm long. Inflorescences with dense grey to rusty, + woolly 
tomentum with hairs 0).2-0.4 mm long, in CO’: 2-3 times ramified, many-flowered, 
(1.5-) 3-11 x 1-6 cm, common peduncie 2-20 mm; in Q: 1.5-7 cm long; bracts 1-2.5 
mm long, caducous. Flowers solitary or in loose clusters of 2-4, perianths 2- (rarely 
up to 4-) valved, glabrescent late and usually towards the base with some persistent 
tomentum of stellate hairs c. 0.2 mm long; pedicel slender, thinly pubescent, at 
base inarticulate. Male perianth subglobose, little to rather much laterally com- 
pressed (subcircular in outline), about as long as broad, 1.5-2 X 1.5-2.1 mm, upper 
and basal parts rounded; pedicel 0.8-2(-2.5) mm long; perianth at anthesis cleft to 
c. Yio, valves 0.2(-0.4) mm thick. Androecium + flattened, in lateral view rounded 
above, c. 0.8-1.2 x 1-1.3 mm; anthers (8-)10-12, suberect, mutually appressed, not 
septate, free apices up to 0.1(-0.2) mm long; androphore up to 0.2 mm long; anther 
column at apex narrowly hollowed for c. “4-3. Female perianth ellipsoid or ovoid 
to obovoid, 2-2.5 xX 1.5-2 mm, split at anthesis to c. %, valves c. 0.4 mm thick, 
pedicel 0.5-3 mm long; ovary ovoid-ellipsoid, densely pubescent with hairs c. 0.1 
mm, 1.2-1.5 « 0.8-1.2 mm, stigma sessile, 2-lobulate, c. 0.3 mm high. Fruits 1-8 per 
infructescence, ellipsoid, apex rounded, base rounded and usually short-contracted 
into the stalk, 1.7-2.5(-3.4) « 1.5-1.8(-2.0) cm, glabrescent but always with minute 
hairs c. 0.1 mm at base (lens!), drying orange-brown or brown, without or with few 
scattered minute tubercles; dry valves 1-2 mm thick; stalk 5-8 mm long; perianth 
not persisting. 


Distribution. Solomon Isls. 


SOLOMON ISLS: Arifanata 2572; BSIP 427, 803, 970, 1124, 1332, 1405, 1537, 1848, 2273, 2582, 
2811, 3035, 3052, 3218, 3318, 3406, 3481, 3679, 3745, 4045, 4046, 4096, 4230, 4834, 5539, 5569, 5617, 
5905, 6230, 6724, 6787, 6900, 7565, 8166, 8362, 8455, 8659, 8916, 9090, 9224, 9427, 9558, 9697, 9955, 
10205, 10256, 10574, 10802, 10978, 11179, 11237, 11408, 11586, 11618, 11714, 12264, 12383, 12523, 
12637, 12782, 13038, 13063, 13235, 13442, 13520, 13360, 13777, 13982, 14088, 14126, 14368, 14475, 
15611, 15860, 15864, 15924 (p.p.), 15967, 16033, 16294, 16364, 16604, 16907, 17370, 17494, 18497, 
18694, 18842; Chapman 427; Hunt 2164; Kajewski 2022; NGF. 31097, 31370, 45611; Waterhouse 891-B: 
Whitmore 6112. 


Ecology. Primary and secondary forest, on a variety of soils; alluvial (sandy, 
clayey) soil, marshy soil, limestone, red soil, ultrabasic and igneous rock; on 
well-drained as well as on (periodically) flooded and marshy ground; not in man- 
grove; 0-850 m. Flowers and fruits throughout the year. 


Vernacular name. Aininiu (Kwara’ae). 


New account of Horsfieldia 2 115 


NOTES 


1. Fieldnotes. Tree usually recorded as without buttresses, but low buttresses 
were noted for BSJP 9090 Guadalcanal). Flowers pale, i.e., pale greenish, pale 
yellow, whitish-yellow, or ivory, strongly sweet scented. Fruits greenish, when fully 
ripe possibly a deeper orange. 


Chapman BSIP 427 (New Georgia) has exceptionally rather chartaceous leaves. 


Fruits measure exceptionally as large as 3.0-3.4 cm, e.g., in BSIP 970, 9427 
(Guadalcanal), BSIP 15967 (Tetepari Isl.). 


2. Related species. Sinclair (l.c.) extensively comments on the relationship and 
postulates its possible hybrid origin from the other two Solomon Isls. species H. 
irya and H. spicata (only p.p., is in my present treatment as H. tuberculata). In my 
opinion, however, the present species is particularly related to H. laevigata (which 
Sinclair erroneously included in H. parviflora), a wide-spread variable species 
which is ‘replaced’ by H. whitmorei in the Solomon Isls.; for differences see the key 
to the species. Besides H. laevigata, the species seems also particularly closely 
related to H. psilantha, under which further notes are presented. 


35. Horsfieldia laevigata (Bl.) Warb. Fig 1B(35); 17 a-i. 


Horsfieldia laevigata (B|.) Warb., Mon. Myrist. (1897) 351, tab. 21, fig. 1-2 (excl. spec. Java) — 
Myristica laevigata B|., Rumphia (1837) 191, t. 64, fig. 3, anal. 1-4; A. DC., Prod. 14, 1 (1856) 202): 
Migq., Fl. Ind. Bat. 1(2), 1 (1858) 65, p.p. — Type: (cult. Mauritius, 2) Commerson 238 (L; iso: P). 


For further synonyms see under the varieties. 


Tree 4-25 m. Twigs terete, faintly ridged or not, towards the apex 1.5-5(-9) m 
diam., early glabrescent, tomentum grey to brown, with hairs c. 0.1-0.2 mm; bark 
striate, when older not flaking; lenticels conspicuous or not. Leaves in 2 rows, 
membranous or thin-chartaceous, elliptic to oblong-lanceolate, broadest usually at 
or above the middle, 10-30 x (3-)4-12 cm, base attenuate, top acute-acuminate); 
upper surface drying dull olivaceous to dark brown, usually minutely paler pustu- 
late, lower surface (largely) glabrescent, the hairs 0.1-0.2 mm, without larger 
brown dots, the nerves not or little contrasting in colour; midrib above flattish;: 
nerves (10-) 12-30 pairs, above thin and flattish or slightly raised, beneath with 
marginal arches usually not very regular and faint; tertiary veins forming a lax 
network, faint or distinct but thin on both surfaces; petioles 5-15 x 1.5-3 mm; leaf 
bud 10-15 x 1.5-2 mm, with hairs 0.1-0.2 mm. Inflorescences subglabrescent or 
with rather dense to sparse scale-like stellate hairs 0.1-0.2(-0.5) mm; in C&: 2-4 
times ramified, many-flowered, 5-20 x 3-10 cm, common peduncle 10-40 mm; in Q: 
c. 2-10 cm long; bracts 2-3 mm long, caducous. Flowers generally in loose clusters 
of 2-5 each; perianths 2- valved, sparsely to densely pubescent (densest towards the 
base) with hairs c. 0.1-0.2 mm long; pedicel not tapering, pubescent, at base not 
articulated. Male perianth as seen laterally subcircular (or sometimes slightly 
longer than broad, or rarely broader than long, e.g., in certain specimens from the 
Papuan Isls.; see notes), usually distinctly flattened, 1.7-2.8 (-3.0) X 1.7-3.0(-3.3) 
mm, upper and basal part rounded; pedicel slender, 1.5-3(-4) mm long; perianth at 
anthesis cleft to 12-% (-%, in certain specimens from the Papua Isls; see notes), 
valves (0.1-) 0.2-0.3 mm thick. Androecium laterally flattened, subquadrangular to 
+ reniform in outline, above broadly rounded to subtruncate, 1.1-1.5 x 1.1-1.8(- 
2.2) mm; anthers 9-16, usually distinctly septate, erect, free portions at apex up to 


Fig. 16. 


116 


Horsfieldia whitmorei Sinclair 

a, twig portion with male inflorescences, X 2; b, male inflorescences, lower down on the 
same twig, X 12; c, mature male flower, x 12; d, ditto, opened, showing androecium, X 12; e, 
portion of twig with female inflorescence, x 2; f, female flower longitudinally opened, 
showing pubescent ovary, X 12; g, portion of twig with infructescence with mature fruits, x 
¥Y2. — a-d, from BSIP 16033; e, from BSIP 3035; f, from BSIP 13442; g, from BSIP 15611. 


ars p iia” 


New account of Horsfieldia 2 117 


0.2 or 0.4-0.6 (var. novobritannica) mm long, the anther column at apex narrowly 
hollowed for c. Y4(-¥2) or in var. novobritannica for c. “10; androphore up to 
0.1(-0.2) mm, + broadly attached. Female perianth broadly ellipsoid to globose, 
2.5-3.1 X 2.8-3.1 mm, split at anthesis to c. %-%, valves 0.3-0.5(-0.8) mm thick, 
pedicel 2-2.5 mm long; ovary ovoid or subglobose 2.0-2.3 xX 1.7-2.2 mm, pubes- 
cent with hairs c. 0.1 mm long or less, style up to 0.3 mm long, stigma sessile, 
minute, hardly bilobulate, c. 0.1-0.2 mm. Fruits (1-)2-15 per infructescence, ellip- 
soid or rarely nearly globose, apex rounded to acutish, base rounded, (1.6-)1.8- 
2.8(-3.0) x 1.4-2.0(-2.2) cm, glabrescent but always with minute hairs c. 0.1 mm 
long at least at base (lens!), drying blackish or greyish-brown, usually with coarse, 
paler coloured tubercles or lenticels; dry valves 2-3 mm or 4-6 mm thick as in some 
forms from SW. New Guinea and New Britain; seed ellipsoid; stalk 3-6 mm long; 
perianth not persisting. 


Distribution. Moluccas, New Guinea, Bismarck Arch. (see further under the 
Varieties). 


A variable, complex species, of which one prominent form is segregated here as 
a Variety. 


KEY TO THE VARIETIES 


la. Hairs on inflorescences 0.1-0.2(0.3) mm long, sometimes almost absent. Anthers at apex free for 
only c. 0.1-0.2 mm: stamen column hollowed for c. 14(-'4, or slightly deeper). Fruit generally 
ellipsoid, pericarp 2-3 mm thick when dry, rarely (SW. New Guinea) 4-6 mm thick. Moluccas, New 
ES Ee @ er ay Ee ee ey a. Var. laevigata 


b. Hairs of inflorescences more woolly, c. 0.3-0.5 mm long. Anthers at apex free for c. 0.4-0.6 mm: 
the column hollowed for c. “%o. Fruit generally subglobose or short-ellipsoid, pericarp 2-5 mm 
Bo, Bh ion GRY ak an wntlad & be ia geKpo non Soanfinwe-anconbar b. var. novobritannica 


a. var. laevigata Fig. 1B(35); 17 a-e 


Myristica nesophila Mig., Ann. Mus. Bot. Lugd. B t. 1(2) (1864) 206, p.p. (excl. sp. from Batjan) — 
Horsfieldia nesophila (Miq.) Warb., Mon. Myrist. (1897) 281, t. 21 fig. 1-2 — Type: Ceram, de Vriese 
s.n. (CO) (L, lecto). 


Horsfieldia polyantha Warb., Mon. Myrist. (1897) 281, t. 23 fig. 1-2; Sinclair, Gard. Bull. Sing. 28 
(1975) 95 (for the greater part, incl. type) — M. polyantha Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 
(1900) 85 — Type: (Aru Isls., Wokam) Beccari s.n. (Acc. Nos. 7619, 7619-A) (FI, n.v.). 


Twigs in apical portion 2-5(-9) mm diam. Leaves 10-30 x 4-12 cm. Inflorescences 
with rather dense to sparse tomentum of hairs 0.1-0.2(-0.3) mm, sometimes almost 
glabrous. Male perianth 1.7-3.3 mm diam.., at anthesis split to c. ¥2-7/4(-%). Anthers 
(9-)10-16, free at apex for (0-)0.1-0.2 mm; anther column at apex hollowed for c. 
Ya(-¥2, or slightly deeper). Infructescence up to c. 10 cm long. Fruits ellipsoid, 
18-28 mm long, drying blackish or brown, usually with coarse wart-like lenticels; 
pericarp 2-3(-6) mm thick. 


Distribution. As the species (including New Britain). 


MOLUCCAS. Commerson s.n. (Bourbon, 238 cult. Mauritius) — Halmaheira: Jdjan & Mochtar 
191; Pleyte 141, 377, 409 (p.p.); de Vogel 3437, 3498 — Bacan (Batjan): de Vogel 3694 — Ceram: b.b. 
25845; Kornassi (exp. Rutten) 996; de Vriese s.n. — Aru Isls (Wokam): Beccari s.n. (FI Acc. Nrs. 7619, 
7619A); Buwalda 5015. 


Fig. 17. Horsfieldia laevigata (BI.) Warb. var. laevigata: a, twig apex with leaves, X 2; b, opened 
mature male flower, showing androecium, X 6; c, androecium, longitudinal section, schema- 
tic, X 12; d, portion of twig with female inflorescence, x 1%; e, female flower, opened, 
showing finely pubescent ovary and minute 2-lobed stigma, x 6. — Horsfieldia laevigata var. 
novobritannica (Sinclair) de Wilde: f, portion of twig with male inflorescence, x 12; g, opened 
mature male flower, showing androecium, x 12; h, androecium, longitudinal section, schema- 
tic X 12; i, infructescence with mature fruits, x 2. — Horsfieldia pilifera Markgraf: j, opened 
male flower, showing androecium, X 6. —a-c, from LAE 52086; d & e, from Commerson s.n. 


(Ile de France) (type); f-h, from Floyd 6430; i, from White NGF 10811; j, from Ledermann 
6675. 


118 


New account of Horsfieldia 2 119 


NEW GUINEA. Irian Jaya: Aet (exp. Lundquist) 348; b.b. 30563, 30595; Branderhorst 165; BW 
(Koster) 1003, 1022, 7149, (Schram) 14943; Doctors van Leeuwen 10623; Pleyte 688; von Rémer 329; 
Soegeng 284 — Papua New Guinea: Brass 1220, 24203; Craven & Schodde 874; Hartley 10147, 10707; 
Jacobs 9242; Kanis 1199; LAE 51683, 51841, 52030, 59069; NGF 2413, 2934, 7275, 14800, 17752, 19625, 
20792, 24080, 28016, 42997, 46543, 48466, 48476; Rau 173; (Vinas) UPNG 3528 — Papuan Islands 
(Fergusson, Normanby, Woodlark): Brass 28660; LAE 52577, 68761, 68871 — Bismarck Arch. (incl. 
Admiralty Isls): LAE 51188, 52086, 52128, 53736, 63063, 66701; NGF 7039, 12980, 26659, 26785, 27303, 
46030, 49511. 


Ecology. Primary and secondary rain forest on ridges and plains, riverine forest, 
swampy scrub and forest, edges of sago-swamps; on a great variety of soils, incl. 
black volcanic soil (Moluccas); 0-1000 m alt. Flowers and fruits throughout the 
year. 


Vernacular names. Kawok-kawoe (Noemfoer Isl.), Kamojer (Noemfoer Isl.), 
Luhakon (Halmaheira), Peita (Western Prov., Papua, Oriomo dial.), Samgoot 
(Kebar lang., Vogelkop). 


Uses. Fruits reported as edible; wood used for house construction (wood reported 
several times as of medium weight and hardness). 


NOTES 


1. Fieldnotes. Bole straight, without buttresses; bark often shallowly vertically 
fissured, not peeling off; branches horizontal, or drooping. Wood whitish or straw. 
Flowers (greenish or brownish yellow. Fruits yellow to orange, aril bright orange. 


2. Variation. After the separation of var. novobritannica the remaining type- 
variety is still a very variable entity. The most marked variations are in the 
following features: 


a. Leaf texture. The leaves are generally membranous, but certain specimens, 
especially some from New Britain, have rather leathery leaves so that the 
tertiary venation may be quite obscure. 


b. Length of hairs of the tomentum on the leaf bud. Hairs in specimens from 
NW. New Guinea (incl. Vogelkop Penins.) may be somewhat longer and 
coarser than usual, and may be up to 0.3 mm long. 


c. Twigs are usually terete, and not or hardly ridged in between the insertion of 
the petioles. A few specimens, e.g., NGF 2430 (in fruit) from N. Prov. Papua, 
or LAE 51683, from Morobe Dist., have rather distinctly lined twigs; such 
specimens may be confused with species with typically ridged twigs, e.g., H. 
iriana or H. angularis. 


d. The size and shape of the mature male perianth. Apparenttly all diameter 
sizes between 1.7-3(-3.3) mm can be found. Specimens of close affinity, with 
still smaller flowers, have been accommodated in the separate species H. 
pilifera. The shape of the perianth in lateral view is generally subcircular. 
Certain specimens, e.g., from the Papuan Isls and Gulf Dist. may have the 
perianth rather markedly broader than long, and these flowers may resemble 
those of, e.g., H. spicata. In specimens from the Moluccas the perianth is 
often slightly longer than broad. 


120 Gard. Bull. Sing. 38(1) (1985) 


e. The degree to which the male perianth opens at anthesis. In most specimens 
the perianth opens to c. 2-way to “3 deep; specimens from the Papuan Isls. 
(e.g., Brass 28660, LAE 52577, 68871), or the Bismarck Arch. (e.g., LAE 
53736, 63063) may have male perianths split at anthesis to as deep as c. %5-%; 
this feature is reminiscent also of H. spicata. In these broad, deeply-splitting 
male flowers, the androecium is relatively broad, and rather reniform as seen 
laterally. 


f. Number of anthers. This varies normally from 12 to 16. Some specimens 
deviate in having a relatively small androecium with apparently only 10(-12) 
anthers; e.g., in Pleyte 141, and other material from the Moluccas. 


g. The thickness of the pericarp. A few specimens, especially from SW. New 
Guinea, e.g., b.b. 30563, 30595, Aet 348, have thick corky pericarps, 4-6 mm 
thick; these specimens perfectly agree in their leaves with var. laevigata. 


3. In the drawing of the type specimen of Myristica laevigata, Commerson 236, a 
female specimen, the perianths are erroneously depicted as 3-valved by Blume in 
Rumphia 1; in the original specimen in L, from which the drawing was obviously 
made, all flowers have 2-valved perianths. 


4. Some forms with larger and thick-walled fruits appear difficult to separate 
from H. pachycarpa; see the notes under that species. 


5. Of Jacobs 9242, from E. New Guinea, mature fruits are in spirit. They are 
apparently similar in size to those in the fresh state, measuring c. 4 x 3.5 cm; the 
perianth at one side of the fruit (up to 15 mm) is much thicker than at the opposite 
side. On drying, these fruits had shrunk to c. 2.5 X 2 cm, with the pericarp only c. 5 
mm thick, thus falling well within the sizes in dry material measured for var. 
laevigata. 


b. var. novobritannica (Sinclair) de Wilde, comb. nov. | Fig. 17 f-i 


Horsfieldia hellwigii var. novobritannica Sinclair, Gard. Bull. Sing. 28 (1975) 54 — Type: Floyd NGF 
6430 (LAE, n.v.; iso: L, K; A, BRI, CANB, NSW, n.v.) 


Horsfieldia novae-lauenburgiae Warb., Mon. Myrist. (1897) 278; K. Schum., Fl, Neu-Pomm. in Notizbl. 
Bot. Gart. Berlin 2 (1898) 117; K Schum. & Lauterbach, Fl. Deutsch. Schutzgeb. Siidsee (1900) 324; 
Markgraf, Bot. Jahrb. 67, 2 (1935) 151 — Type: Bismarck Arch., Neu Lauenburg Group, Ulu Isls., 
Warburg 20713 (B +; G, iso, n.v.; identity not sure, see notes). 


Horstieldia ralunensis auct. non Warb., Kanehira & Hatusima, Bot. Mag. Tokyo 52 (1938) 355 
(specimen Kanehira 3969 n.v.). 


Twigs in apical portion 2-6(-8) mm diam. Leaves (12-)17-30 x (3-) 5-8 cm. 
Inflorescences with rather dense woolly tomentum of hairs c. 0.3-0.5 mm long. 
Male perianth c. 1.8-2.0 x 2.2 mm, at anthesis split to c. “%. Anthers c. 14, almost 
completely filling the perianth, septate, at apex free for c. 0.4-0.6 mm; anther 
column from the apex hollow for c. “o. Infructescences up to c. 8 cm long. Fruits 
broadly ellipsoid to almost globose, 18-22 x 16-20 mm, drying grey-brown, with 
coarse wart-like lenticels; pericarp rather hard, 2-3(-5 mm, see notes) thick. 


Distribution. Bismarck Arch.: New Britain. 


NEW BRITAIN: Floyd (NGF) 6430, 6662; NGF 10040, 10811, 21731, 25504, 32622. 


New account of Horsfieldia 2 121 


Ecology. Primary and disturbed rain forest; 0-1000 m. Flowers in August, fruits 
throughout the year. 


Vernacular names. La gele kuku (W. Nakaina), Nungan (S. New Britain). 
NOTES 
1. Fieldnotes. Fruits globose, golden brown, yellow, or orange at maturity. 


2. Relationship. The var. novobritannica, of which the male flowers are only 
known from the type, deviates within H. laevigata by its androecium. This com- 
pletely fills the perianth; at the apex the anthers are mutually free to almost 
halfway, and the column is hollow from the apex to c. “10 deep. The column is 
reminiscent of that in H. irya, and probably the variety originated by some 
hybridization with the latter. In this respect it can be mentioned that the inflor- 
escences of H. irya from this region may be similarly woolly hairy, and also that the 
leaves of the type of the var. novobritannica show whitish markings, as found 
regularly in H. irya. 


3. Vegetatively, the fruiting specimens resemble strongly the flowering type 
specimen, but one can not be quite sure whether all belong to the var. novobritan- 
nica. The fruits of Floyd 6662 rather deviate as they are almost globose, c. 22 mm 
diam., and have a thick spongy pericarp c. 5 mm thick; this condition is possibly 
pathological. 


4. On the label of Floyd 6430, the type, is commented that it is almost the same 
species as Floyd 6410. This is not so, as Floyd 6410 is a good H. hellwigii var. 
hellwigit. 


5. The identity of H. novae-lauenburgiae Warb. is not clear to me. The holotype 
Warburg 20713 was burnt in B, but Sinclair (p. 116) has seen an istotype in G, 
enumerated under his broad conception of H. spicata. He has not commented on 
this specimen. I have not seen this isotype. 


According to the original description the specimen is large-leaved, with rather 
persistent tomentum, the female inflorescences much-branched, the flowers 2- 
valved, + pubescent, the ovary tomentose, and this may well point to Sinclair’s H. 
hellwiggi var. novobritannica, the basionym of the present new combination. In 
Markgraf’s opinion (p. 151) H. novae-lauenburgiae is close to H. hellwiggii. 


36. Horsfieldia pilifera Mkef. Fig. 1B(36); 17 j 
Horsfieldia pilifera Markgraf, Bot. Jahrb. 67, 2 (1935) 154 — Type: Ledermann (B, 7; iso: L) 


Horsfieldia novoguineensis Warb., Mon. Myrist. (1897) 271, p.p., Hollrung 657, syntype; lectotype= H. 
aruana. 


Tree (5-)10-20 m. Twigs terete, lined or faintly ridged or not, towards apex 
1.5-4(-10) mm diam., early glabrescent, tomentum grey to brown, of hairs c. 
0.1 mm; bark striate, when older not flaking; lenticels usually inconspicuous. 
Leaves in 2 rows, membranous, elliptic to oblong-lanceolate, broadest at about the 
middle, 7-27 x 2.5-8.5 cm, base attenuate, tip acute-acuminate; upper surface 
drying brown, usually minutely whitish pustulate, lower surface early glabrescent, 


122 Gard. Bull. Sing. 38(1) (1985) 


hairs stellate, 0.1 mm long, without larger brown dots, the nerves not particularly 
contrasting in colour; midrib flat or slightly raised above; nerves 7-16 pairs, above 
thin and flat, beneath with the marginal arches faint and not very regular; tertiary 
veins forming a lax network, faint on both surfaces; petioles 6-12 x 1.5-2.5 mm; 
leaf bud c. 10 X 1-2 mm, with hairs c. 0.1 mm long. Inflorescences sparsely to 
densely pubescent with rather woolly hairs 0.1-0.3 mm; in CO: 2-4 times ramified, 
many-flowered, 5-12 x 4-8 cm, common peduncle 10-25 mm, in Q: 4-12 cm long; 
bracts 1-2(-3) mm long, caducous. Flowers generally 2-5 together; perianth 2- 
valved, sparsely to densely pubescent with stellate hairs c. 0.1 mm long; pedicel 
slender, not tapering, finely pubescent, at base inarticulate. Male perianth, as seen 
laterally, subcircular or sometimes slightly transversely or longitudinally elliptic, 
laterally little to much flattened, (1.0-)1.2-1.8 x 1.2-1.8(1.9) mm, upper part 
broadly rounded, at base rounded to short-attenuate, pedicel 1-2 mm long; 
perianth at anthesis split to 13-2, valves c. 0.1 mm thick. Androecium flattened, 
subquadrangular in outline, above broadly rounded, 0.7-1.2 < 0.6-1.1 mm; anthers 
8-10, (sub)erect, septate, free portions at apex up to 0.1 mm long, the anther- 
bearing column at the top narrowly hollowed for c. %-'%; androphore up to 
0.1 mm, broadly attached. Female perianth broadly ellipsoid c. 2.8 X 2.5 mm, cleft 
at anthesis to 2-%4, valves 0.5-0.8 mm thick, pedicel c. 2 mm long; ovary globose to 
ovoid, c. 1.5 X 1.3 mm, pubescent with hairs c. 0.1 mm or less, style and stigmas 
minute, c. 0.1 mm long. Fruits (2-)5-20 per infructescence, globose to short- 
ellipsoid, top and base rounded, 1.1-1.6 x 1.1-1.6 cm, glabrescent but always with 
minute hairs persistent towards the base (lens!), drying bright to dark brown, 
without or with little larger paler tubercles; dry valves often somewhat woody, 
thickest at one side, 1-3 mm thick; seed ellipsoid; stalk 1-5 mm long; perianth not 
persisting. 


Distribution. Northern half of New Guinea: Vogelkop, Japen Isl., Jayapura, 
Sepik, Madang, Morobe Prov. 


NEW GUINEA. Irian Jaya: bb. 30429, 30547, 30558; Brass 14014; BW (Koster) 1097, (Versteegh) 
4811, (Schram) 6082, (Kalkman) 6245, (Versteegh & Vink) 8298 — Papua New Guinea: Clemens 524, 
635, 1710, 10825; Hartley T.G.H 11029; Hollrung 657; Hoogland 5139, (& Craven) 10275; Jacobs 9609, 
9609A; LAE 52756, 73818; Ledermann 6675, 10450; NGF 28080; Schlechter 16933. 


Ecology. Primary and secondary rain forest; reported from sandy loam soil, 
mixed forest with Anisoptera at c. 100 m; 0-1000 m alt. Flowers and fruits through- 
out the year. Fruiting once reported as very prolific. 


Vernacular names. Gaben (Morobe Dist.), Gefrah (Tehid lang., W. Vogelkop), 
Guma (Waskuk lang., Sepik Prov.), Mamgananieproi (Biak lang). 


NOTES 


1. Fieldnotes. Bark longitudinally fissured; sap watery, turning pink or red; wood 
straw to brown, of moderate weight and hardness. Flowers yellow. Fruits hard, 
glossy green, turning dark yellow, orange or red, aril orange-red. 


2. Relationship. A species very close to H. laevigata, distinguished by the smaller 
male perianth, and smaller globose or subglobose fruits with or without but little 
coarser, paler-coloured lenticel-like tubercles. The male perianths rather vary in 
outline e.g., those of Ledermann 6675 being rather lengthwise ellipsoid and c. 1.5 
x 1.3 mm whereas those of Ledermann 10450 (type) and Clemens 1710 are + 
transversely ellipsoid, measuring c. 1.2 X 1.6 mm and 1.5-1.8 xX 1.5-1.9 mm 
respectively. 


New account of Horsfieldia 2 123 


—~_ 


The female flowers of Schlechter 16933(K) only have been seen and described by 
me; the perianth is generally somewhat smaller than in H. laevigata, but the female 
flowers probably do not differ significantly in the two species. 


Because of the fruits, which are sometimes globose, the present species may be 
confused with H. basifissa, one which has quite different male flowers. 


3. Sinclair (p. 112) included the present species in H. spicata var. spicata, a very 
large taxon as conceived by him. 


37. Horsfieldia lancifolia de Wilde, sp. nov. Fig. 1B(37); 18 


Horsfieldia species foliis lanceolatis, 5-16 cm longis. perianthio masculo pynformi, 2.5-3 mm longo, 
sparse pubescenti, antheris, 6-8, erectis, ovario pubescenti, fructibus ellipsoideis vel pynformibus. 
2.5-3.5 cm longis in siccitate, glabrescentibus. — Type: b.b. Cel./II- 464 (L; iso: K: BO & SING, n.v.). 


Tree, 10-30 m. Twigs terete, towards the apex 1-2.5(-5) mm diam., not or but 
faintly llined, glabrescent from tomentum composed of hairs 0.1 mm or less; bark 
finely striate, when older not flaking; lenticels smallish, not very conspicuous. 
Leaves in 2 rows, (thinly) chartaceous, oblong-lanceolate to lanceolate, broadest at 
or somewhat above the middle, 5-16 x 1.5-3.5 (-4.5) cm, base (long-)cuneate, tip 
acute-acuminate; upper surface drying olivaceous to brown, with or without small 
whitish marks (without larger irregularly-shaped whitish marks), lower surface 
early or late glabrescent, hairs c. 0.1 mm or less, provided or not with brownish 
dots and points of mixed sizes; midrib above flattish or slightly raised; nerves 9-17 
pairs, thin, flat, and inconspicuous above, the submarginal arches + regular in 
shape but indistinct; tertiary venation forming a rather lax network, inconspicuous; 
petiole slender, 10-20 x 1-1.5 mm; leaf bud 10-20 x 1.5-2 mm, with hairs c. 0.1 
mm. Inflorescences with rather sparse tomentum of hairs 0.1-0.2 mm long, in C: 2 
or 3 times ramified, not many-flowered, c. 5 x 3.5 cm, common peduncle c. 10 mm 
long, flowers solitary or 2 or 3 together; 2 inflorescences short, 1-3 cm long, 
slightly ramified, 4-10-flowered; perianths 2-valved, rather sparsely pubescent with 
hairs 0.1 mm or less in length, pedicels pubescent through hairs 0.1-0.2 mm, at base 
inarticulate. Male perianth obovoid to pear-shaped, not much laterally compress- 
ed, drying blackish, not or little collapsed, 2.5-3 x 2.0-2.3 mm, above broadly 
rounded, at base tapering into the pedicel c. 1.5-2 mm long; perianth at anthesis 
cleft only for c. 1/6, valves (at base) thickish, c. 0.4-0.5 mm thick. Androecium 
(synandrium) long-obovoid, laterally flattened, 1.5-1.8 x 1.0-1.2 mm, above sub- 
truncate; anthers 6-8, erect, free apices c. 0.3-0.4 mm: anther column solid, except 
for the apical portion in-between the free apices of the anthers; androphore rather 
slender, c. 0.6-0.8 mm long. Female perianth subglobose to obovoid, variable in 
size (see notes), 2-3 < (1.8-)2-3.5 mm, at anthesis cleft to c. ¥4-%, valves (perianth) 
c. 0.4-0.8 mm thick, pedicel 1-2 or 3-4 mm long, pubescent with hairs 0.1-0.2 mm; 
ovary ovoid, 1.4-2.2 x 1.0-2.0 mm, densely minutely pubescent; style and stigma(s) 
minute, c. 0.1 mm. Fruits 1-4 per infructescence, ellipsoid, (and + contracted at 
base) or pear-shaped (see notes), 2.5-3.5 xX 1.8-2.4 cm, completely glabrescent. 
granulate or with small lenticels or tubercles, drying brown: dry valves thick- 
woody, 4-8 mm thick; seed broadly ellipsoid, c. 17 x 14 mm; stalk 3-10 mm long: 
perianth not persisting. 


Distribution. Central and South Celebes. 


Fig. 18. 


124 


Horsfieldia lancifolia de Wilde 


a, habit of leafy twig with male inflorescences, x 12; b, mature male flower, lateral view, x 6; 
c, ditto, opened, showing androecium, X 6; d, androecium, longitudinal section, schematic, x 
6; e, portion of twig with female axillary inflorescences, leaves fallen, x 2; f, mature female 
flower, lateral view, < 6; g, ditto, opened, showing finely pubescent ovary, x 6; h, portion of 


twig with infructescences with mature fruit, x 2. — a-d, from b.b. Cel./II-464 (type); e-g, 
from de Vogel 5267; h, from van Balgooy 3973. 


New account of Horsfieldia 2 125 


CELEBES. Central: van Balgooy 3142; de Vogel 5267 — South (-Central): b.b. Cel./IJ-236, -404, 
-464; van Balgooy 3931, 3973, 4083; de Vogel 6243, 6287. 


Ecology. Forest on ultrabasic rock (iron, nickel), laterite, also limestone ridges; 
200-1200 m alt. Flowers throughout the year; fruits from May to July. 


Vernacular name. Tabenoe benoe (Malili area). 


NOTES 


1. Fieldnotes. Flowers buds brown; fruits glossy green, turning yellow-green to 
orange. 


2. The Q-flowering specimens de Vogel 5267, van Balgooy 3142 (from Mt. 
Roroka Timbu, Central Celbes, 1100-1200 m alt.), and the one in fruit, de Vogel 
6243 (S. Celebes, N. of Lake Matano, c. 400 m.), differ from the rest of the 
material from both Central and S. Celebes by, respectively, much stouter flowers 
with larger 9 perianths (3-3.5 mm diam.) and larger fruits, subellipsoid in shape, c. 
3.5 cm long. In the other specimens the Q perianths are only c. 2-2.5 mm, and the 
fruits generally more pear-shaped, c. 2.5-3.0 cm long. 


3. Sinclair identified the then-known specimens, including the type of the present 
new species, as H. parviflora (Roxb.) Sinclair. The new species is easily recognized 
among Celebes-Moluccan material by its pear-shaped male flowers, its largish 
fruits with thick pericarp, and its small, rather narrow leaves. The androphore is 
proportionally long as compared to other species. 


38. Horsfieldia decalvata de Wilde, sp. nov. Fig. 1B(38) 


Horsfieldia species floribus bivalvatis, H. tuberculatae atque H. laevigatae similis, ab eis differt 
floribus masculis breviter pyriformibus, perianthio tenuiter pubescenti, androcio compresso, antheris 6 
apice non-incurvis, ovario pubescenti, fructibus subglobosis c. 1-1.2 cm diam., tenuiter pubescentibus. 
— Type: Idjan & Mochtar 181 (L; iso: BO, n.v.; K). 


Tree 10-15 m. Twigs subterete, towards the apex + flattened and faintly ridged 
or not, 1.5-4(-9) mm diam., early glabrescent, tomentum brown, composed of 
hairs c. 0.1 mm; bark dull brown, finely striate, when older not flaking, lenticels 
small, not very conspicuous. Leaves in 2 rows, membranous, elliptic-oblong to 
oblong-lanceolate, 11-25 x 3.5-7 cm, base attenuate to long-attenuate, tip acute- 
acuminate; upper surface drying dark brown, lower surface early glabrescent, 
hairs c. 0.1 mm; without brown dots; midrib above flat; nerves c. 12-16 pairs, above 
thin and flattish, on lower surface with the marginal arches not very distinct; 
tertiary venation on upper surface as a fine network, + distinct or not; petioles c. 
5-10 x 1.5-2.5 mm; leaf bud slender, c. 6-10 x 1-2 mm, densely dull brown 
pubescent by hairs c. 0.1 mm long or less. Inflorescences c. 2(or 3) times ramified, 
in CO: c. 4-6 X (1-)2-4 cm, in 9 up to c. 2.5 cm long; branches subglabrescent 
tomentum minute with hairs c. 0.1 mm; common peduncle in male 1-2 cm; bracts 
and bracteoles not seen, caducous. Flowers in C& solitary or 2 or 3 together, 
minutely pubescent through hairs c. 0.1 mm long or less; perianths 2-valved. Male 
perianth subglobose to short-pear shaped, moderately laterally compressed, as long 
as it is wide, c. 2.3 x 2.3 mm, upper part broadly rounded, the lower half tapering 
into the tapering pedicel c. 1.5-2 mm long, short-pubescent, inarticulate at base; 


126 Gard. Bull. Sing. 38(1) (1985) 


perianth at anthesis cleft to c. 73, valves c. 0.2-0.3 mm thick. Androecium mod- 
erately flattened, in outline c. 1.6 X 1.4 mm, upper part rounded; anthers c.6 (i.e., 
c. 12 thecae with the connectives rather broad), not septate, c. 1.6 mm long, erect, 
not inflexed, free apical parts up to c. 0.1 mm; androphore up to c. 0.1 mm long; 
anther column cleft to c. 4-4. Female flowers not seen; immature fruits (ovaries) 
densely finely pubescent. Fruits c. 2-5 per infructescence, short-ellipsoid to subglo- 
bose, c. 1.1-1.2 x 1.0-1.1 cm, minutely pubescent with hairs c. 0.1 mm or less, 
drying brown, with scattered small tubercles; dry valves c. 1 mm thick; stalk 
slender, c 4 mm long; perianth not persisting. 


Distribution. Moluccas. 


MOLUCCAS. Morotai: Kostermans 767. — Halmaheira: Idjan & Mochtar 181. — Ceram: Buwalda 
5627. — Ambon: Robinson 1878. 


Ecology. Forest at low altitudes, 0-100 m. Flowers in September, fruits from May 
to November. 


Vernacular name. Sekukumailor (Halmaheira). 
NOTES 
1. Fieldnotes. Flowers recorded as brown, fruits yellow. 


2. H. decalvata superficially resembles very much a number of species including 
wide-spread species like H. moluccana, H. tuberculata and H. laevigata, but our 
present new species is distinct by its finely pubescent, pear-shaped male flowers, 
erect anthers, pubescent ovaries, and finely pubescent, small subglobose fruits. H. 
tuberculata also has pear-shaped flowers which are generally glabrous, and it has 
larger, glabrous fruits. H. /aevigata has the male perianth more spherical in outline, 
and the fruits much larger and pubescent. H. moluccana has incurved anthers and 
glabrous fruits. See also note 3. 


3. In 1975 Sinclair determined the specimens (now assigned to the present 
species by me) as H. parviflora, a species, which in my present treatment differs in 
general habitat (pale twigs), quite different spike-like inflorescences (glabrous male 
flowers with different androecium; glabrous ovaries) and in its glabrous, larger 
fruits, which blacken on drying. In his description of H. parviflora, he erroneously 
accepted both the glabrous and the tomentulose condition of the ovaries for that 
species. 


39. Horsfieldia tuberculata (K. Sch.) Warb. Fig. 1B(39); 19 


Horsfieldia tuberculata (K. Sch) Warb., Mon. Myrist. (1897) 279, t. 23 f. 1-3; K. Sch., Notizbl. Bot. 
Gart. Berl. 2 (1898) 117; Schum. & Lauterbach, Fl. Deutsch. Schutzgeb. Siidsee (1900) 324; 
Markgraf, Bot. Jahrb. 67, 2 (1935) 151, p.p.; A.C. Smith, J. Arn. Arb. 22, 1 (1941) 62. — Myristica 
_tuberculata K. Sch. in Schum. & Hollrung, Fl.. Kais.-Wilh. Land (1899) 46; Warb., Bot. Jahrb. 13, 
3-4 (1891) 308. — Type: (Bat Isl. Admiralty Isls.) Hollrung 848 (O’, Kaiser Wilhelms-Land) (B 7+; iso: 
K, L & P); Kdrnbach s.n. (fr., B Tt). 


H. novoguineensis var. moseleyana Warb., Mon. Myrist. (1897) 273; K. Schum. & Lauterbach, FI. 
Deutsch. Schutzgeb. Stidsee (1900) 324. — Type: Moseley s.n. (B +; iso: BM, K). 


H. solomonensis A.C. Smith, J. Arn. Arb. 22, 1 (1941) 64. — Type: Kajewski 1549 (A, n.v.; iso: BM, P; 
BO, BRI & G, n.v.). 


Fig. 19. 


Horsfieldia tuberculata (K.Sch.) Warb. 

a, habit of leafy twig with infructescence, x 1; b, portion of twig with male inflorescences, x 
Y2; c, mature male flower, lateral view, x 6; d, ditto, opened, showing androecium, X 6; e, 
portion of twig with female inflorescence, x 12; f & g, opened female flower and glabrous 
ovary with shallowly 2-lobed stigma, x 6 and xX 12 respectively; h, twig portion with 


infructescence with mature fruits, x 2.—a, from BSIP 14035; b-d, from Waterhouse 820-B; 
e-g, from BSIP 9628; h, from BSIP 10611. 


127 


128 Gard. Bull. Sing. 38(1) (1985) 


Tree 5-20 m. Twigs terete, faintly ridged or not, towards apex 2-4(-6) mm diam., 
early glabrescent, tomentum (rarely + woolly), of hairs 0.1-0.3 mm, bark 
striate, when older not flaking, lenticels sparse and small, or sometimes almost 
absent. Leaves in 2 rows, membranous (or from higher altitudes sub-chartaceous), 
elliptic to oblong-lanceolate, 12-25(-40) x 3-10(-16) cm, base short- to long- 
attenuate, or rarely rounded, tip acute-acuminate; upper surface drying dull 
greenish brown to brown, often minutely pustulate, lower surface early glabres- 
cent, hairs 0.1(-0.3) mm, without dark brown dots; midrib above flat or + sunken; 
nerves 11-22 pairs, above thin and flattish or sunken, beneath with the marginal 
arches not very distinct; tertiary veins forming a rather fine network distinct or not 
on both surfaces; petiole 8-15(-18) x 1.5-3(-5) mm; leaf bud 10-15 x 2-3 mm, with 
hairs 0.1-0.3 mm long. Inflorescences with sparse tomentum of stellate-dendroid 
hairs 0.1-0.3 mm long, sometimes + glabrescent, in O 1-3 times ramified, 3-15 
(-22, see notes) X 2-10 cm, common peduncle 5-25 mm long; in 9: up to 7 cm long; 
bracts 0.5-3 mm long, caducous. Flowers solitary or in loose clusters of 2-4, 
glabrous (early glabrescent); perianths 2-(or 3-) valved; pedicels glabrous or rarely 
sparingly pubescent (NGF 169, New Britain), at base not articulated. Male 
perianth as seen laterally short-pear shaped, laterally compressed, generally about 
as long as broad to slightly broader than long, (1.5-)2-3.5 x 2-4 mm, upper part 
broadly rounded, in the lower 3-2 usually narrowed into the somewhat tapering 
pedicel 2-5(-6) mm (pedicel sometimes only little tapered, e.g. Brass 21765); 
perianth at anthesis split to 2-7/4, valves c. 0.2 mm thick. Androecium + flattened, 
broadly rounded to subtruncate above, 1.5-2.5 x 2-3 mm; anthers (12-)14-20, 
faintly septate, 1.5-2.5 mm long, erect, not inflexed, free apical parts 0-0.2 mm; 
anther column at apex narrowly hollowed for “5-3; androphore up to 0.2 mm long. 
Female perianths subglobose, 2-3 xX 2-3.5 mm, split at anthesis to 12-%, valves 
0.4-0.8 mm thick; pedicel 1.5-2.5 mm long, ovary subglobose, glabrous, 1.5-2 x 
1.5-2 mm; stigma sessile, faintly 2-lobed, c. 0.2 x 0.5-1 mm. Fruits 5-15 per 
infructescence, ellipsoid, apex rounded or slightly pointed, 1.5-3.7 X 1.1-2.5(-3.0) 
cm, glabrous, drying blackish brown, with scattered or sparse, rather small to 
coarse, paler, lenticel-like tubercles; dry valves 1-8 mm thick; stalk c. 3-10 mm; 
perianth not persisting. 


Large-fruited specimens from the Papuan Isls. Dist. are accommodated in a 
separate variety. 


KEY TO THE VARIETIES 
la. Fruits 15-25 x 11-16 mm. Dry pericarp 1-2(-3) mm thick ................cccceeeeeeeeees a var. tuberculata. 


b. Fruits27-37 x 17-25 mm: Dry pericarp 3-3.mm ticks) .............4ih «02 eee b. var. crassivalva 
a. var. tuberculata 


Distribution: Caroline Isls. (Palau Isls.), Admiralty Jsls., Bismarck Arch., Solo- 
mon Isls., Papuan Isls., New Guinea: New Guinea: Cape Vogel Penins. 


CAROLINE ISLS.: Tuyama 9349; Masahiko Takamatsu 526. 
ADMIRALTY ISLS.: LAE 53629, Hollrung 848; Moseley s.n. 


BISMARCK ARCH.: Commerson s.n.; LAE 52150, 66532; NGF 169, 7944, 10981, 12329, 26574, 
26621, 26737, 29680, 29683, 32227, 41370, 4140, 42280; Sands et al. 2975. 


PAPUA NEW GUINEA (mainland): Brass 21765; Saunders 493; UPNG. 4035. 


Wow. aWerm *- 


New account of Horsfieldia 2 129 


PAPUAN ISLANDS: Brass 28464; LAE 52607, 74575, NGF 25280. 


SOLOMON ISLS.: BSIP 426, 737, 766, 1125, 1273, 1967, 2248, 2615, 2855, 3145, 3358, 3437, 3510, 
3703, 3739, 3843, 3986, 4154, 4183, 4719, 4842, 4873, 4960, 4998, 5180, 5396, 5519, 5743, 5806, 5922, 
5689, 5997, 6172, 6391, 6414, 6849, 6858, 7110, 7186, 7419, 7717, 7736, 7826, 8164, 8395, 8721, 8880, 
S912, 9046, 9165, 9323, 9628, 9783, 9951, 10057, 10179, 10223, 10295, 10393, 10522, 10576, 10611, 
10697, 10721, 10824, 10887, 10913, 10952, 11374, 11284, 11766, 12212, 12565, 12651, 12836, 12864, 
12917, 12958, 12992, 13066, 13131, 13281, 13347, 13402, 13558, 13682, 13798, 13871, 13948, 14025, 
14035, 14078, 14099, 14127, 14202, 14205, 14342, 14488, 14553, 14625, 14826, 14868, 14900, 14929, 
14981, 15074, 15161, 15260, 15281, 15556, 15568, 15871, 16000, 16141, 16422, 16507, 16595, 16769, 
16784, 16895, 16915, 16982, 17436, 17538, 17802, 17880, 17987, 18045, 18221, 18313, 18478, 18683, 
18722, 18758, 18835, 18912, 19421; Brass 2605, 2983, 3460; Craven & Schodde 90; Hunt 2387; Kdrnbach 
1892, sterile (L); Kajewski 74(75), 1549, 2554, 2710; McKinnon 7; NGF 16356, 16391, 16431, 19709, 
25280, 31120, 45632, 45693, 45753; Schodde (& Craven) 3687, 4068; Waterhouse 35 (21275), 167, 178, 
820-B: Whitmore 6022, 6188, 6290. 


Ecology. Primary and secondary forest; on coral rock, seashores, limestone, 
swamp forest; 0-700 m. alt. Flowers and fruits throughout the year. 


Vernacular names. Abuino’o, Ambuino’o, Ambuynor (Kwara’ae lang, Solomon 
Isls.); Aininiu, Ainynu (Kwara’ae lang., Solomon Isls.); Kokotetepina (Kwara’ae 


lang., Solomon Isls.); Bale bale (New Britain); Pive’ar (Mbuke lang., Manus 
Prov.). 


On the label of a Whitmore specimen from Choiseul Isl. is written that 
‘Ambuynor’ differs from ‘Aininiw’ (= H. whitmorei) in the shorter, broader leaves. 


NOTES 


1. Fieldnotes. Flowers several times recorded as yellow, sweet scented or with 
strong smell. Fruits yellow, orange or orange-brown. Exudate of bark red, watery. 
Slash wood white or brownish white, soft; slash bark soft, pale brown or reddish 
brown. 


2. Variability. The sizes of flowers and fruits are available; for instance, the 
perianths of NGF 45632 (C’), Waterhouse 820-B (©), both from Bougainville Isl., 
BSIP 9628 (Q, Rennell Isl.), and Brass 2983 (Q Ulawa Isl.) measure 3-3.5(-4) mm 


diam., whereas Whitmore BSIP 3843 (C’) and 737 (9), from Guadalcanal measure 
only c. 2 mm diam. 


3. The leaves of specimens from Bat Isl. (Admiralty Isls.), incl. the type of H. 
tuberculata, can reach up to c. 40 by 14 cm. 


The inflorescences of the type of H. tuberculata are as large as 12-18 cm long, 
those of Commerson (port Prastin) s.n. (P.) reach 22 cm. 


The pedicels of the male flowers in Brass 21765 (Cape Vogel Penins.) are 
slender, and only slightly tapering; such specimens may easily key out wrongly. 
They probably represent a separate taxon as yet insufficiently defined. 


Also, tab. 23, fig. 1, drawn by Warburg, was obviously made from the type in B; 
it does not show the perianths and pedicel as being much tapered but the latter is 
rather terete and slender. 


On the other hand, I have examined the istotypes in K, L, and P and found them 
agreeing well with the description of the species as presently given. 


130 Gard. Bull. Sing. 38(1) (1985) 


b. var. crassivalva de Wilde, var. nov. 


A varietate typica in fructibus maioribus 27-37 mm longis pericarpio sicco 3-8 mm crasso differt — 
Type: Brass 28352 (L: K. iso). 


Twigs rather stout, at apex (3-)4-S mm diam. Leaf blades 25-35 x 10-12.5 cm. 
Male perianths c. 3 xX 3.3 mm; androecium much flattened, c. 1.8 X 1.8 mm, 
anthers c. 12, free apices c. 0.1 mm long; androphore c. 0.1 mm long. Pedicels 3-6 
mm. Fruits 27-37 X 17-25(-30) mm. Dry pericarp c. 3-8 mm thick. 


Distribution. New Guinea, Louisiade Arch.: Misima Isl., Tagula Isl., Rossel Isl.: 
doubtful on San Cristobal (Solomon Isls.). 


LOUISIADE ARCH.: Brass 27648, 28143, 28352; Whitmore 6268 (San Cristobal, doubtful). 


Ecology. Riverine rain forest at low altitudes, creek alluvium soil; 0-20 m. 


NOTES 


1 Fieldnotes. Subcanopy tree. Flowers yellow, very fragrant. Fruits to 5 cm 
diam., orange, ovoid or subglobose, keeled; aril pink. 


2. Specimens belonging to the present variety generally have a stout habit with 
coarse twigs and large leaves, and have relatively large perianths; these sizes fall, 
however, within those accepted for the type variety. 


3. The fruits of the type, Brass 28352, from Rossel Isl., at sea-level, are in the 
fresh state reported as subglobose and as large as 5 cm diam. Those of Brass 27648 
(Misima Isl.) measure c. 4.5 X 3.5 cm. A specimen from Rossel Isl. included in var. 
tuberculata, Brass 28464, has mature fruits, which when dried are only c. 18-22 mm. 
This was collected at c. 700 m alt. 


4. The specimen Whitmore 6268, from San Cristobal is doubtful; it has stout 
leaves, the dry fruits measure c. 27 mm long, but the pericarp is only c. 2 mm thick. 
It probably belongs to var. tuberculata. 


5. Specimens in fruit may be confused with H. pachycarpa, but this has minutely 
pubescent fruits (and pubescent ovaries), at least at the base towards the insertion 
of the stipe. 


40. Horsfieldia corrugata Foreman Fig. 1B(40); 20 a-c 


Horsfieldia corrugata Foreman, Contr. Herb. Austr. 10 (1974) 45, fig. 1. — Type: Foreman & Lelean 
LAE 5246] (LAE, n.v.; iso: K, L; BRI, CANB, A, E & SING, n.v.). 


Tree 5-12 m. Twigs terete, not ridged, towards the apex (3-)4-5(-12) mm diam., 
early glabrescent, tomentum minute greyish to rusty, of hairs c. 0.1 mm; bark 
striate, when older not flaking, lenticels present, coarse but usually not much 
contrasting in colour. Leaves in 2 rows, thinly coriaceous, elliptic-oblong, broadest 
at or above the middle, 12-29(-32) x 4.5-8.5(-10) cm, base attenuate, top acute- 
acuminate; upper surface drying dark brown, minutely pustulate or not, lower 
surface early glabrescent, without larger dark-brown dots; midrib slender to rather 
broad, flattish above; nerves 12-18 pairs, thin and flat above, beneath with the 


“= aVier 


Fig. 20. 


ie 


Rs, 
= 
= 
~, 
. 
= 


Horsfieldia corrugata Foreman: a, longitudinally opened male flower showing androecium, X 
6; b, ditto, female flower, showing pubescent ovary and narrow 2-lobed style, x 6; c, fruit, x 
Y2. — Horsfieldia pachycarpa A.C. Smith: d, leafy twig with infructescence, x '/2; e, longitudi- 
nally opened male flower, showing androecium, X 6; f, ditto, female flower with pubescent 
ovary with shortly 2-lobed stigma, X 6; g, almost mature fruit, x ’2.— a, from Carr 14123; b, 
from LAE 60020; c, from Carr 14334; d, from LAE 62196; e, from LAE 51940; f, from 
Clemens 5378; g, from NGF 38895. 


131 


132 Gard. Bull. Sing. 38(1) (1985) 


marginal arches distinct or not, not very regularly looping; tertiary veins forming a 
lax network, indistinct; petiole 6-18 x 2-3.5 mm; leaf bud 10-20 x 1.5-3 mm, with 
hairs c. 0.1 mm. Inflorescences thinly pubescent with rusty stellate hairs c. 0.1 mm 
long or less; in CG’: 2 or 3 times ramified, rather slender, (4-)6-14 x 2-9 cm, in 9 up 
to c. 5 cm long, common peduncle c. 10 mm; bracts pubescent, 1.5-4 mm long, 
caducous. Flowers (in ©’) solitary or in loose clusters of 2-5, glabrous or glabrescent 
from scattered hairs less than 0.1 mm; perianth 2-valved; pedicel + tapering, thinly 
pubescent or glabrescent, at base inarticulate. Male perianths in lateral view 
subcircular, including the short-pear shaped pedicel, which is somewhat laterally 
compressed, about as broad as long, 3.0-3.5 x 3.0(-4.0) mm, the top broadly 
rounded, the lower half + tapering into the thickish tapering pedicel (2-)3-4 mm 
long; perianth at anthesis splitting from 12 to nearly ~4, valves c. 0.2 mm thick at 
apex, the perianth towards base only slightly thicker, often provided with a few 
coarse blackish-brown wart-like dots. Androecium thickish, not much laterally 
compressed, above broadly rounded, (1.5-)2.0-2.2 x 2.0-2.2(-3.0) mm, anthers 
8-12, erect, not septate, c. 2 mm jong, free apical parts 0.1-0.2 mm, androphore 
Q.2-0.3 mm long; anther column narrowly hollowed for c. “s-'4 at apex. Female 
perianth rather narrowly ovoid, almost glabrous, with a few coarse, dark-brown 
wart-like dots, c. 4.5 x 3 mm, cleft at anthesis to c. %4-!4, valves c. 0.3-0.4 mm 
thick, coriaceous; pedicel 4-5 mm long, very minutely scattered-pubescent; ovary 
ovoid, somewhat dented or corrugated, c. 2.5-3 x 2.5 mm, densely minutely 
pubescent with hairs less than 0.1 mm long, style and 2-lobed stigma glabrous, 
0.8-1.0 mm long. Fruits 1(-4) per infructescence, ramiflorous, broadly ellipsoid to 
subglobose, somewhat flattened, 6-7.5 * 4.5-6.5 cm, at base contracted or not into 
a short pseudostalk, top acutish, coarsely flanged and corrugated, drying blackish 
brown, with scattered coarse paler-coloured tubercles, glabrescent, valves + 
woody-corky, 10-20 mm thick; stalk 5-10 mm long; perianth not persisting. 


Distribution. Papua New Guinea: Central Prov., Northern Prov., Milne Bay 
Prov. 


NEW GUINEA. Papua New Guinea: Carr 14123, 14334, Foreman & Lelean LAE 52461, 60020; 
NGF 41016, 46430; Pullen 5496. 


Ecology. Primary and secondary rainforest on slopes and ridges, fagaceous 
forest; at 1200-1900 m. Flowers and fruits from July to December. 


Vernacular name. Sodowa (Port Moresby Dist.) 
NOTES 

1. Fieldnotes. Mountainous terrain; small tree; wood very light brown. Flowers 
yellow or orange. Fruits green, strongly wrinkled or corrugated, and strongly 


ridged. Aril orange. 


2. I have not seen the male flowering material described by Foreman, but the 
male flowers of Carr 14123, not cited by him, agree with his description. 


3. Those specimens of this species, not seen by Foreman, were identified by 
Sinclair as H. spicata var. spicata (Carr 14334) or H. praetermissa Sinclair, in sched. 
(Carr 14123). ; 


New account of Horsfieldia 2 133 


4. When in flower, the specimens may be difficult to distinguish from, e.g., H. 
pachycarpa or H. tuberculata. Possibly the few coarse and conspicuous blackish- 
brown wart-like dots on the perianth, found in both CG and Q flowers, are 
characteristic for the species. Furthermore, the very large corrugated and ridged 
thick-valved fruits are very distinctive. Large, thick-valved fruits also occur in H. 
pachycarpa, H. tuberculata var. crassivalva or in certain forms of H. laevigata, but 
in Our present species the fruits exceed those in size. 


41. Horsfieldia pachycarpa A.C. Smith Fig. 1B(41); 20 d-g 
Horsfieldia pachycarpa A.C. Smith, J. Arn. Arb. 22 1 (1941) 64 — Type: Brass 610 (A, n.v.). 


Horsfielda praetermissa Sinclair, in sched. (Carr 13262, etc.). 


Tree; 5-25 m. Twigs terete, faintly ridged or not, towards the apex 3-5(-12) mm 
diam., early glabrescent, tomentum minute, with hairs c. 0.1 mm; bark coarsely 
striate, when older not flaking, lenticels usually coarse and distinct. Leaves in 2 
rows, membranous or thinly coriaceous, elliptic-oblong to lanceolate, broadest 
below to above the middle, 17-30 x 4-11 cm, base attenuate, tip acute-acuminate; 
upper surface drying dark brown, usually minutely pustulate, lower surface early 
glabrescent, without larger dark-brown dots; midrib above slender to rather broad, 
flattish; nerves 14-18 pairs, above thin, flat, inconspicuous, beneath with the 
marginal arches distinct or not, not very regularly looping; tertiary veins forming a 
lax network, indistinct; petiole 6-12 X 2-5 mm, leaf bud 10-15 x 2-3 mm, with hairs 
c. 0.1 mm. Inflorescences subglabrous or with sparse stellate hairs 0.1-0.2(-0.3) 
mm, 2-4 times ramified, in C’: 7-15 x 6-10 cm, in 2 up toc. 10 X 4.cm, common 
peduncle 5-25 mm; bracts caducous, not seen. Flowers in loose clusters up to 5 
each, thinly pubescent with hairs c. 0.1 mm, or glabrescent towards the apex; 
perianth 2-valved; pedicels thinly pubescent, at base not articulated. Male 
perianths short-pear shaped, laterally somewhat compressed, about as broad as 
long or slightly longer than broad, 2.5-3.2 x 2.8-3.2 mm, upper part bluntish to 
broadly rounded, the lower *%4 more or less tapering into the thickish and much 
tapering pedicel 2-5 mm long; perianth at anthesis cleft to nearly '2, valves towards 
apex c. 0.2 mm thick, the perianth towards base usually thicker, + firm-fleshy or 
coriaceous, 0.3-0.8 mm thick. Androecium + laterally flattened, c. 1.8-2.1 x 
1.1-1.8 mm, above broadly rounded, anthers 5 or 6, or 9 or 10, erect, not or hardly 
septate, c. 1.5-2 mm long, free apical parts c. 0.1-0.2 mm; androphore rather 
slender, 0.2-0.5 mm.; anther column at apex narrowly hollowed for c. %-'%. 
Female perianths ovoid-ellipsoid, 2.5-4 x 2.5-3.2 mm, split at anthesis to c. 4, 
valves 0.4-0.6 mm thick; pedicel 2-4 mm long; ovary ovoid, densely minutely 
pubescent, 2.5-3 x 1.8-2.5 mm, stigma sessile, minutely 2-lobed, c. 0.1 mm. Fruits 
1-6 per infructescence, ellipsoid to broadly ellipsoid, often + ridged towards the 
base, rounded or tapering into a short pseudostalk, apex rounded, (3.0-)3.5-4.5 x 
(1.8-) 2.0-3.0 cm, minutely pubescent towards the base, or glabrescent, drying 
blackish(-brown), usually with conspicuous coarse, paler-coloured lenticel-like 
tubercles; dry valves + woody, (4-)5-10 mm thick; stalk 5-10 mm; perianth not 
persisting. 


Distribution. New Guinea. West (Irian Jaya): Snow Mountains (Idenburg R., 
1300 m); East: West Sepik Prov., Western Highlands, Eastern Highlands, South- 
ern Highlands, Madang, Morobe, Central Provinces. 


134 Gard. Bull. Sing. 38(1) (1985) 


NEW GUINEA. West (Irian Jaya): Brass 610, 12752. — Papua New Guinea: Carr 13262 (13362), 
14146; Clemens 5378; H.O. Forbes (7) 192; LAE 51940, 60363, 60365, 62196, 65747; Manner & Street 
307; NGF 21297, 29213, 29357, 32652, 38895, 47888. 


For deviating specimens see the notes. 


Ecology. Primary and secondary montane forest, forest clearings, etc.; often on 
ridges; recorded from fagaceous and Castanopsis-Lithocarpus forest; (450-)1000- 
2000 m alt. Flowers and fruits throughout the year. 


Vernacular names. Cheeweng (Maring), Kupgne (West Sepik), Mong-mong 
(Weng lang., West Sepik). 


NOTES 


1. Fieldnotes. A montane species; branches often horizontal. Bark shallowly 
vertically fissured; exudate watery, clear or + reddish. Wood pink, cream, or 
whitish. Flowers yellow, fragrant. Fruits ramiflorous, glossy green turning yellow to 
orange, eaten by cus-cus. 


2. Related and resembling species. Differs from H. tuberculata by the more 
thick-fleshy or woody-fleshy, largely pubescent perianth, and by the pubescent 
ovary and fruit. 


Fruiting specimens may resemble H. laevigata much, which is obviously closely 
related. The male flowering specimens of the present species (Carr 13262, 13362, 
14146, LAE 51940, Manner & Street 307) have, however, essentially differently 
shaped (+ pear-shaped) flowers, and are of a more fleshy-woody consistency; those 
of H. laevigata being much more globose, with the pedicel more slender, not 
pear-shaped, and of a more membranous-herbaceous consistency. 


Through the somewhat resembling male flowers H. pachycarpa appears related 
to H. corrugata, a species with much larger ‘corrugated’ fruits, from similar mon- 
tane habitats. The flowers of H. corrugata are probably always provided with some 
large, thickened blackish dots, a character not seen in H. pachycarpa. 


It should be remarked here that H. tuberculata var. crassivalva (from the 
Louisiade Arch.), a taxon only known from fruits which also have a thick pericarp, 
is very similar as well. 


3. Deviating specimens. Rather many specimens in fruit deviate in the smaller 
sizes of female flowers and fruits, viz. Carr 13901, Jacobs 8834, 9079, LAE 66790, 
NGF 37317, 41594, Robbins 624. They are from Central Prov., W., E, and S. 
Highlands Prov., and West Sepik Prov.; all from well above 1000 m., except Jacobs 
9079 (600-700 m alt.). The female perianths measure about 2-2.8 x 1.8-2.3 mm; the 
fruits c. 2.5-3.5 x 1.7-2.0 cm, and have a thick woody pericarp. In size and general 
appearance these fruiting specimens seem intermediate to and often can hardly be 
distinguished from the common H. laevigata. However, the fruits of the mentioned 
specimens also completely link up with the generally larger fruits of H. pachycarpa, 
of which the male flowers are presumably essentially of a different shape and 
texture as those of H. laevigata. Future collectors should search in the field for male 
specimens which belong with certainty to the above-mentioned female collections. 


TiN 


New account of Horsfieldia 2 135 


4. Sinclair included most specimens of the present H. pachycarpa in his H. 
spicata var. spicata; Car 14146 was identified provisionally as H. praetermissa 
Sinclair, in sched., with the remark that better material is still required. 


5. I have not seen the type specimen, Brass 610 (A) from Biriatabu at 450 m 
(Snow Mts. Dist.) but its description agrees well with the species as described 
presently. In P there is an isotype, without fruit. 


42. Horsfieldia pulverulenta Warb. Fig. 1B(42) 


Horsfieldia pulverulenta Warb., Mon. Myrist. (1897) 342, t. 23 fig. 1-2; Markgraf, Bot. Jahrb. 67, 2 
(1935) 150 (sub H. ralumensis) — Myristica pulverulenta (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 
1 (1900) 87 — Horsfieldia hellwigii var. pulverulenta (Warb.) Sinclair, Gard. Bull. Sing. 28 (1975) 56 
p.p., excl. Vink BW. 12194 = H. leptantha — Type: NW. New Guinea, Vogelkop Penins. (Andai, 
Mt. Arfak), Beccari 759 (FI, n.v.), 925 (FI, n.v.). 


Horsfieldia hellwigi var. hellwigii x var. pulverulenta (Warb.) Sinclair, Gard. Bull. Sing. 28 (1975) 58 — 
Syntype: Saunders 202 (L, lecto), 358, 398, 483 (L: CANB, n.v.). 


Tree 15-25 m. Twigs terete, towards the top 4-10 mm diam., early to rather late 
glabrescent, tomentum dark rusty with hairs c. 0.5-1.2 mm, bark finely striate or 
not, when older not flaking, lenticels usually present. Leaves in 2 rows, coriaceous, 
elliptic to oblong-lanceolate, broadest usually at or below the middle, or + parallel- 
sided, 14-35(-40) x 4-10.5(-13) cm, base rounded or short-attenuate, tip acute 
acuminate, often to 2(-3 in sapling shoots) cm, caudate; upper surface drying usually 
dull brown to olivaceous, minutely rugose-pustulate, lower surface late glabrescent 
or with persistent tomentum composed of + evenly sized and spaced, rather harsh, 
dark brown hairs c. 1.0-1.5 mm long, when shed usually leaving thickened and rough 
hair bases, without brown dots; midrib above flattish, later glabrescent and usually 
with persistent tomentum towards base; nerves 11-30 pairs, generally rather 
straight, 50-70° with the midrib, c. 5-15 mm apart, thin and sunk above; beneath the 
marginal nerve with the arches usually very regular and prominent; tertiary vena- 
tion rather lax, above well-visible and sunken, giving the blade often a + bullate 
appearance; petiole 2-12 x 2.5-4.5 mm, not or hardly winged; leaf but stout, 3-6 cm 
long, with harsh hairs 1-1.5 mm long. Inflorescences woolly-pubescent with hairs 
1-1.5 mm long, 2-3 times ramified, rather many-flowered, in Q and GC’: 4-10 x 2-9 
cm, common peduncle up to 15 long; bracts (broadly) ovate, acutish, 3-5 mm long, 
caducous. Flowers + solitary (in Q) or in loose clusters of 2-6; perianths 2-valved, 
largely set with stellate(-dendroid) hairs 0.1-0.3 mm; pedicels slender, pubescent 
with coarse hairs 0.4-0.7 mm, at base inarticulate. Male perianths in lateral view 
subcircular, 1.5-3 x 3-4 mm, the basal part thick and coriaceous, the remainder 
collapsed when dry and perianth then often saucer-shaped or wrinked, at the apex 
just above the anthers, opening by a minute pore-like slit less than 1 mm wide; 
valves or apical part of perianth c. 0.2(-0.4) mm thick. Pedicels 2-3.5 mm long. 
Androecium consisting of a coriaceous + ellipsoid column c. 0.8-1.1 mm long, with 
2 small anthers, each 0.2(-0.3) mm at the apex. Female perianth broadly ellipsoid- 
ovoid, 3.8-4 xX 3.5-4 mm, split at anthesis to c. %-Y10, with a minute pore-like slit 
above the stigmas, valves c. 0.3-0.5 mm thick; pedicels (1.5-)3-5 mm long; ovary 
ovoid-subglobose, c. 2.5-3 X 2.5 mm, densely pubescent with hairs 0.1-0.3 mm; 
style erect, glabrous, 0.2-0.8 mm long; stigma 2-lobed, c. 0.2-0.3 mm long. Fruits c. 
3-10 per infructescence, ellipsoid, top acute and sometimes acuminate, 3.0-5.0 x 
2.0-3.0 cm, minutely pubescent at least at the base, valves woody-coriaceous, 4-7 
mm thick, usually with paler, small or coarse lenticels or tubercles; stalk 2-7 mm 
long; perianth not persisting. 


136 Gard. Bull. Sing. 38(1) (1985) 


Distribution. New Guinea: Irian Jaya (Vogelkop, Jayapura, Geelvink Bay; 
Mimika, in the South), Papua New Guinea (West Sepik, East Sepik, Madang, 
Western; the Gulf Prov. specimen deviates. 


NEW GUINEA. Irian Jaya: b.b. 30514, 31098, (Lundquist 103) 32822; BW 2727, 2733, 6666, 9173, 
9420, 11111; Doctors van Leeuwen 10479 — Papua New Guinea: Darbyshire & Hoogland 8094; Katik W 
2877; LAE 52930, 53567; NGF 27471, 34154, 34339, 45861, 45913, 48290; Saunders 202, 358, 398 — 
Deviating: Gulf Dist., Schodde & Craven 4662 (see notes). 


Ecology. Lowland primary and (old) secondary rain forest, ridge-side forest, 
swamp forest; on clay, stony-sandy soil; 0-500 m alt.; flowers and fruits throughout 
the year. 


Vernacular names. Baa (Hollandia), Gumaga (E. Sepik), Ibuumkwaraf (Kem- 
toek lang., Hollandia), Patepa (Siere-Octa, SW. New Guinea), Poi (Pogatumo 
lang., Sepik Prov.), Sabobo (Orne lang., Mafoka, Sepik Prov.), Vionge (Nemo 
lang., Hollandia). 


NOTES 


1. Fieldnotes. Bole unbuttressed or with slight buttresses. Bark often strongly 
peeling in small, oblong, thin scales, black-brown or dark brown. Wood whitish or 
straw, moderately hard and heavy. Flowers yellow, greenish-yellow or pale orange- 
yellow. Fruit greenish, greenish-yellow, or yellow-brown; aril red. 


2. The present species obviously belongs to the group with H. hellwigii which has 
stout twigs, the pubescence on the leaf bud and apex of twigs conspicuous, com- 
posed of coarse hairs (0.5-)1-1.5 mm long. It is very distinct by its woody perianths, 
usually collapsed in O’ around the much ‘reduced’ androecium. This latter consists 
of an ellipsoid woody body, its apex with only 2 apparently much reduced anthers 
or thecae just below the apical pore-like slit of the perianth. 


The female flowers are larger than those of H. hellwigii. 


3. The late Dr. Muller of the Rijksherbarium investigated the pollen of LAE 
52930 and remarked: little pollen produced, 25-30 yu, boat-shaped with proximal 
side convex, exine 0.5 yw thick, very finely echinate, echinae 0.5 pw long. In 
comparison, the related Horsfieldia hellwigii (Brass 25949) produces abundant 
pollen, 30-40 4, boat-shaped with convex proximal side, exine 1.5 yw thick, finely 
echinate, echinae 1 yw long. 


4. Deviating specimen. The specimen Schodde & Craven 4662, from the Gulf 
Province, Papua, with C flowers, deviates in its non-coriaceous leaves, the more 
woolly tomentum (not harsh) and the densely tomentose perianths. The perianths 
do not dry blackish brown as is usual in H. pulverulenta, but are instead, rather 
light brown and only slightly collapsed. The perianths contain a much reduced 
androecium, comparable to that found in H. pulverulenta. The marginal nerve of 
the leaves is looping very regularly, as in H. pulverulenta. The specimen may 
represent a separate taxon. 


5. Sterile, dubious specimens. The sterile material cited below with rather mem- 
branous leaves, probably belong to the present species. For the greater part they 
might have been taken from sapling shoots. The tomentum is generally softer than 


New account of Horsfieldia 2 137 


in a true H. pulverulenta. Often the leaves are long-caudate. They might belong to 
one of the other related species of the H. hellwigii-complex as well. Sinclair 
referred these specimens to a group of hybrids between H. hellwigii var. hellwigii 
and var. pulverulenta. The specimens are from the Madang district, NE. New 
Guinea: Saunders 202, 358, 398, and from Jayapura, Irian Jaya: BW 6666, 9173. 


6. Sinclair accepted the present species as a variety under H. hellwigii. 


He, however, did not cite the curious woody flowers with the apparently re- 
duced androecium nor the differing female flowers. 


7. | have not seen the syntypes; Sinclair enumerates them among several collec- 
tions also seen by me, some of which, however, e.g., Vink BW 12194 is presently 
referred to a new species, H. leptantha. 


43. Horsfieldia leptantha de Wilde, sp. nov. Fig. 1B(43) 


Ramuli validi, eoram apicibus atque gemmis praeditis pilis 0.5-1.0 mm longis. Perianthium masculum 
2-valve, subglobosum, pubescens, in anthesi usque as 74-*% divisum, valvis c. 0.2 mm crassis, androecii 
lateraliter compresso, antheris 10-14, erectis. Fructus breviter ellipsoidei, 2.0-2.4 cm longo, grosse 
tuberculati — Type: Irian Jaya, Fak-Fak, Vink BW 12194 (L). 


Tree 8-30 m. Twigs terete, towards the apex 4-8 mm diam., early to late 
glabresent, tomentum rusty, of hairs 0.5-1.0 mm long, bark finely or coarsely 
striate, when older not flaking; lenticels usually present. Leaves in 2 rows, membra- 
nous to chartaceous, elliptic to oblong-lanceolate, broadest usually in the middle, 
generally not parallel-sided, 13-35 x 5-13.5 cm, base (short-) attenuate, tip acute- 
acuminate, not caudate; upper surface drying dull brown to olivaceous, smooth or 
finely pustulate, lower surface rather early to late glabrescent or with persistent 
tomentum of stellate-dendroid rather woolly or ‘mealy’ hairs of mixed sizes, 0.5-1.0 
mm long, when shed not leaving rough hair bases; without brown dots; midrib 
above flattish, glabrescent except towards base; nerves 10-20(-30?, see notes) pairs, 
usually at c. 45°-50° with the midrib, 5-20 mm apart, above thin and sunken or flattish, 
below, the arches of the marginal nerve not very regular, prominent or not; 
tertiary venation lax, distinct or not; petiole 5-12 x 2.5-4 mm, not winged; leaf bud 
stoutish, 2.5-5 cm long, with dense tomentum of hairs 0.5-1.0 mm long. Inflore- 
scences woolly-pubescent with hairs 0.5-1.0 mm long, in CG’: rather many-flowered, 
2 or 3 times ramified, c. 11 X 7 cm, common peduncle 5-20 mm long; flowers in 
loose clusters of 2-6 each, perianth 2-valved, completely pubescent with hairs 
0.2-0.3 (-0.5) mm; pedicels slender, pubescent, at base inarticulate; bracts broadly 
elliptic, obtuse, 2-3 mm long. Male perianth in lateral view subcircular, c. 2.5 X 
2.7-3 mm, somewhat laterally compressed, sub-membranous, not collapsing on 
drying; pedicel 1-1.5 mm long; perianth at anthesis split to 4-%; valves c. 0.2 mm 
thick. Androecium laterally compressed, c. 1-1.3 x 1.3-1.6 mm, above broadly 
rounded-truncate, the column inside narrowly hollowed for c. % (c. 0.3 mm), 
anthers 10-14, erect touching each other, somewhat septate when young, 1-1.1 mm 
long, free apices c. 0.2-0.3 mm, not or but faintly incurved; androphore + 0. 
Female flowers not seen. Infructescences 6-12 cm long, branched. Fruits 2-8 per 
infructescence, broadly ellipsoid to subglobose (seed ellipsoid), apex obtuse or 
broadly rounded, 2.0-2.4 x 1.6-2.0 cm, largely glabrescent but with minute vesti- 
gial hairs towards the base, tubercles coarse, pale, lenticel-like, valves woody- 
coriaceous, 4-7 mm thick; stalk 5-10 mm long; perianth not persisting. 


138 Gard. Bull. Sing. 38(1) (1985) 


Distribution. New Guinea: Irian Jaya (Vogelkop, doubtful; Fak-Fak), Papua 
New Guinea (West Sepik). 


Ecology. Secondary and primary forest, ridge forest, on clay soil over limestone; 
0-600 m. Flowering and fruiting apparently not seasonal. 


NEW GUINEA. Irian Jaya: BW (Koster) 10763 (doubtful, see notes), Vink 12194, (Moll) 12952; 
Pleyte 553; Pulle 343. — Papua New Guinea: West Sepik, LAE 52962; NGF 25241. 


NOTES 
1. Fieldnotes. Bark recorded as greenish black; blaze with pinkish red serous sap. 


2. The male perianths are deeply cleft, to c. /4-%, much deeper than in the other 
species of the H. hellwigii complex. The fruits with their coarse lenticels and 
tubercles are strongly reminiscent of those in H. laevigata; there is likely to be a 
true close relationship with this species. 


3. A doubtful sterile specimen from Vogelkop, BW 10763, is tentatively included 
in the present species, mainly because of its locality. It deviates by its numerous 
side-nerves, c. 30-33, which are almost parallel and depart from the midrib at an 
angle of c. 70°; the marginal nerve is distinct and loops very regularly, both 
characters predominant in the related H. pulverulenta. However, the tomentum on 
the lower leaf surface is rather mealy and is composed of hairs of rather mixed 
sizes, and the Jeaves are membranous in texture, not rough. The specimen might 
likewise belong to one of the other related species of the H. hellwigii complex. 


44. Horsfieldia hellwigii (Warb.) Warb. Fig. 1B(44); 21 


Myristica hellwigii Warb., Bot. Jahrb. 18 (1893) 192 — Horsfieldia hellwigii (Warb.) Warb., Mon. 
Myrist. (1897) 343; Markgraf, Bot. Jahrb. 67, 2 (1935) 150; A.C. Smith, J. Arn. Arb. 22, 1 (1941) 61; 
Sinclair, Gard. Bull. Sing. 28 (1975) 49 (for the type variety only) — Type: Finschhafen (Papua New 
Guinea), Hellwig 416 (B, t+, n.v.). 


H. glabrescens Warb. in K. Schum. & Lauterbach, Fl. Deutsch. Schutgeb. Siidsee (1900) 325 — Type: 
Papua New Guinea (Madang), Tappendeck 74 (B, +; WRCL, seen by Sinclair). 


Tree 5-30 m. Twigs stout, terete or rarely + ridged, often hollow, towards apex 
(4-)5-15 mm diam.; early to late glabrescent from rusty woolly or felty tomentum of 
hairs 0.5-1.0(-1.5) mm long; bark finely or coarsely striate, when older not flaking; 
lenticels present, usually distinct. Leaves in 2 rows, membranous or rarely 
chartaceous, elliptic-oblong to oblong-lanceolate, broadest at or above the middle, 
or sometimes + parallel-sided, 17-40(-50) x 5-14 cm, base + rounded to (short-) 
attenuate, tip acute-acuminate (in New Britain specimens rarely short-caudate); 
upper surface drying usually dull olivaceous, minutely pustulate, lower surface 
late glabrescent or with persistent tomentum of hairs of mixed to subequal sizes 
0.3-1.0 mm long, when shed not leaving thickened hair-bases; without brown dots; 
midrib above flat or slightly sunken, late glabrescent or usually with persistent 
tomentum towards base; nerves 12-33 pairs, thin, + flat above, generally at an 
angle of c. 45° with the midrib, the marginal nerve with arches usually rather thin 
and + irregular beneath; tertiary venation lax and thin, faint above; petiole (2-) 5-8 


2. 


New account of Horsfieldia 2 139 


x 2.5-5 mm, not or hardly winged; leaf bud generally stout, 3-7 cm long, densely 
pubescent, hairs 0.5-1.5 mm long. Inflorescences woolly-pubescent, hairs 0.5-1.0 
mm; in CG: 3.5-15 x 2-10 cm, 2-3 times ramified, flowers in clusters of 3-6 each; in 2 
up to c. 8 X 5 cm, flowers in clusters of up to 4; common peduncle up to 30 mm; 
bracts elliptic to broadly ovate, acutish, 3-7 mm long, caducous; perianths 2- valved 
(flowers not known in var. lignosa), glabrous or early glabrescent except at base, 
pedicels pubescent by stellate-dendroid hairs 0.2-0.5 mm, at base inarticulate. Male 
perianths, as seen laterally, subcircular, 1.8-3.2 x 2.3-3.5 mm, somewhat laterally 
compressed, top and base broadly rounded, pedicel slender, 1-3(-4) mm; perianth 
at anthesis split to (14-) 12-%4, valves c. 0.2 mm thick. Androecium (1.2-) 1.5-2 x 
(1.3-) 1.7-2 mm, laterally compressed to c. 0.8-1 mm thick, above + broadly 
rounded-truncate, column c. 0.3-0.5 mm, narrowly hollowed for the upper c. 
Y4(-5), anthers (10-) 12-18 (i.e., 12-18 thecae at each side), erect, not septate, c. 
1.2-2 mm long, completely sessile, mutually touching, free apices 0-0.1 mm, not 
incurved; androphore + broad, up to 0.1 mm long. Female perianth subglobose or 
broadly ovoid, c. 2.8-3.5 x 3 mm, split at anthesis to 3-12, valves 0.2-0.3 mm thick; 
pedicel 1-2 mm long; ovary subglobose, c. 2 mm diam., densely pubescent with 
hairs 0.2-0.4 mm long, stigma largely sessile, faintly 2- lobed, c. 0.1 mm high. Fruits 
3-15 per infructescence, subglobose or broadly ellipsoid to fusiform, top rounded or 
generally acutish (when dry), 1.2-2.8 x 1.0-1.8 cm, densely pubescent or partly 
glabrescent with hairs c. 0.5 mm, usually rather finely lenticellate-tuberculate, dry 
valves 1-3 mm, or in var. lignosa 4-8 mm thick; stalk 1-4 mm long; perianth not 
persisting. 


Distribution. Papua New Guinea, New Britain and New Ireland; not yet found in 
Irian Jaya. 


Ecology. Primary and secondary forest, regrowth; in primary forest an 
understorey or second storey tree, in secondary forest often common; lowland rain 
forest, ridge forest, also monsoon forest, gallery forest; on alluvial soils, also 
limestone; 0-1200 m. Flowers throughout the year, fruits predominantly from July 
to December. 


Vernacular names. Apaap (Wanigela lang.), Camarngur (Lae subdist., Morobe), 
Fohja (Okema lang. Aku), Guma (Waskuk lang., Sepik Dist.), Hota (Garaina 
lang., Bulolo), Lagele Kuku (W. Nakanai, New Britain), Naufora (Talasea, New 
Britain), Mamasoh (Onjob lang., Koreaf), O’hénga (Orakaiva lang., Mumumi). 


Uses. Fruits sometimes recorded as edible. 
NOTES 

1. Fieldnotes: Bole straight, unbuttressed, bark finely longitudinally fissured; 
crown narrow, dense; branches often tending to be whorled, horizontal, later on 


drooping; leaves drooping; wood rather soft and light, whitish or straw, heartwood 
pinkish. Flowers yellow. Fruits green turning yellow or orange, aril orange. 


Fig. 21. 


140 


he 


| 
: 
; 
: 
| 
: 


\ \ | {\ ( WZ 


Horsfieldia hellwigii (Warb.) Warb. var. hellwigii 

a, apical portion of leafy twig, x '; b, portion of twig with male inflorescence, x 1/2; ¢, 
mature male flower, lateral view, x 6; d, ditto, longitudinally opened, showing androecium, 
x 6; e, portion of twig with female inflorescences axillary to leaf-scars, x 12; f, female flower 
at anthesis, X 6; g, ditto, longitudinally opened, showing pubescent ovary and small bi-lipped 
stigma, X 6;h, infructescence with mature fruits. — a, from NGF 26412; b-d, from NGF 4019; 
e-g, from Hoogland 3431; h, from LAE 67101. 


New account of Horsfieldia 2 141 


2. Taxonomy. The present species, here comprising three varieties, is largely 
identical with Sinclair's H. hellwigii var. hellwigii. Sinclair's var. pulverulenta 
(Warb.) and his var. pulverulenta, Vink BW 12104, are here referred to different 
species. Also, H. ralunensis, which Sinclair cited in the synonymy of H. hellwigii 
var. hellwigii, is presently accepted as a separate species. His var. novobritannica is 
presently partly, almost exclusively the type, referred to a variety of H. laevigata. 


Horsfieldia hellwigii, as presently accepted, is still rather variable, especially in 
fruit size and shape; for some very small-fruited specimens and for specimens with 
conspicuously woody fruits, separate varieties are accepted. 


KEY TO THE VARIETIES 


la. Fruit slightly asymmetrically subglobose; pericarp woody, 4-8 mm thick, the surface not wrinkled on 
OST 2 ee UR SS 2 PB URE ED 5 ney Se a c. var. lignosa 


b. Fruit subglobose, or ellipsoid, or fusiform; dry pericarp c. 2 mm thick, the surface usually somewhat 


oe bal Se a on AS Sea SoS ES) TRS cep pe Oh ee ee ar eee 2 | 2 

2a. Fruit broadly ellipsoid to nearly globose, 12-15 mm long ............................ b. var. brachycarpa 
b. Fruit broadly ellipsoid to fusiform, 16-28 mm long. ............. 00... e cee ee eee ee scene eee ee ees a. var. hellwigii 
a. Var. hellwigii Fig. 1B(44); 21 


Lower leaf surface usually rather densely pubescent with hairs c. 0.3-1.0 mm. 
Male perianth (in bud) 2.2-3.2 x 2.4-3.5 mm, split at anthesis to 12-73. Anthers 
(12-)14-18. Fruit broadly ellipsoid to fusiform, 1.6-2.8 x 1.0-1.8 cm, pericarp 2-3 
mm thick. 


Distribution. As for the species. 


NEW GUINEA. Papua New Guinea — Bismarck Arch. (New Britain, New Ireland): Floyd 6410; 
Hoogland 3431; LAE 75366; NGF 10018, 10832, 24233, 26253, 26310, 27248; Sands et al. 2013; 
Waterhouse 877 — Main Island (Madang, Morobe, Northern, Milne Bay. Western, Central): Brass 
24386; Carr 11962, 13901; Clemens s.n. (8.11. 1936), 78, 10908; Fallen et al. 282; Floyd & Hoogland 
3813; Frodin UPNG 2139; Fryar 4019; Hartley TGH 9970, 10013; Hoogland 3256; 4386, 5210; LAE 
67101; NGF 204, 857, 4019, 1617, 7331, 9677, 9681, 9770, 10540, 10981, 17129, 24003, 24329, 26412, 
28553, 28776, 31697, 33857, 34154, 34339, 36304, 36773, 37659, 41170, 41421, 41823, 44388, 45861, 
45921, 47431, 48290; Saunders 483; Womersley & Hoogland 5156 — Fergusson Isl. (Milne Bay Prov.): 
Brass 25949; LAE 52558 — Deviating specimens: LAE 66008 (Morobe Prov.): NGF 40599 (W. New 
Britain); see notes. 


NOTES 
1. NGF 24233, from limestone at c: 1200 m, has rather chartaceous leaves. 


2. The twigs are normally terete, and not or faintly ridged, in NGF 37656 the 
twigs are very distinctly ridged in between the bases of the petioles. 


3. Further deviating specimens: NGF 40599 from W. New Britain, and LAE 
66008 from nearby Umboi Isl. rather differ one from the other, the former being 
much more pubescent. Both deviate from normal H. hellwigii var. hellwiggi by the 
rather more pear-shaped, i.e., at base more tapering male perianths. Possibly the 
specimens are a hybrid, e.g., with H. tuberculata, and they may appear to represent 
a separate taxon. 


142 Gard. Bull. Sing. 38(1) (1985) 


b. var. brachycarpa de Wilde, var. nov. 


Differt a var. hellwigii fructibus siccis fere globosis, 12-15 mm longis, pericarpio 2 mm crasso — Type: 
Lauterbach 1191 (L; iso: BRSL & S, n.v.). 


Lower leaf surface rather sparsely hairy, the hairs c. 0.3 mm long. Male perianth 
c. 2 mm diam. (1.9 X 2.3 mm), split at anthesis to c. 43. Anthers 10-12. Fruits 
subglobose to broadly ellipsoid, 1.2-1.5(-1.7) x 1.0-1.3 cm; pericarp 1-2 mm thick. 


Distribution. Papua New Guinea (Sepik, Madang, Morobe Prov.). 


NEW GUINEA. Papua New Guinea (Northern): Hoogland & Craven 10312; Lauterbach 1191; NGF 
9146, 10258. 


Ecology. Twice collected in levee-forest; 0-100 m. alt. 
NOTES 


1. The original description of H. hellwigii was based on Hellwig 416, now lost. In 
his monograph of 1897, p. 344, Warburg mentions Lauterbach 1191, with fruits, as 
a specimen assigned to H. hellwigii. It was accepted later by Markgraf (p. 150) and 
Sinclair (p. 52) under H. hellwigii. Now it serves as type of my present new variety 
brachycarpa. 


2.1 have accepted NGF 9146, with male flowers, as belonging to var. 
brachycarpa. It has slightly smaller perianths and fewer anthers as compared with 
the type-variety. 


c. var lignosa de Wilde, var. nov. 


Differt a var. hellwigii fructibus siccis pericarpio lignoso 4-8 mm crasso — Type: Milne Bay Prov., 
Leach LAE 56060 (L: iso: K; A, BISH, BO, BRI, CANB, SING, SYD, PHN & US, n.v.). 


Lower leaf surface rather sparesely hairy, the hairs c. 0.2-0.3 mm long (on leaf 
buds c. 0.5 mm long). Flowers unknown. Fruits somewhat asymmetrically 
subglobose, slightly flattened or not, c. 1.6-1.9 cm diam (immature); pericarp very 
woody, 4-8 mm thick, the surface not wrinkled on drying, densely rusty tomentose 
with hairs c. 0.3(0.5) mm long. 


Distribution. SE. Papua New Guinea: Central and Milne Bay Provinces. 


Ecology. Lowland and mountainous forest, 300-1150 m alt. Fruits in June and 
September. 


NOTES 


J. One suspects the immature fruits in both cited collections as being diseased, 
e.g., infected by a gall, but on closer inspection all fruits seem healthy. The fruits of 
both collections are recorded as green. 


2. The specimen NGF 32401 has the lower leaf surfaces finely, iregularly, more 
darkly mottled; mottles are absent in the type. In the NGF 32401 the older twigs 
are rather markedly ridged in-between the petiole scars. 


New account of Horsfieldia 2 . 143 


3. The specimens cited were collected after Sinclair’s revision Horsfieldia. Flow- 
ers are unknown; when these get collected, it may turn out that the present taxon 


can better be regarded as a separate species. Vegetatively it is very like the typical 
H. hellwigit. 


45. Horsfieldia ralunensis Warb. Fig. 1B(45) 


Horsfieldia ralunensis Warb., Mon. Myrist. (1897) 336; K. Schum.. Notizbl. Bot. Gart. Berl.-Dahl. 
(1898) 117: Sch. & Laut., Fl. Deutsch. Schutzgeb. Siidsee (1900) 324); Markgraf. Bot. Jahrb. 67, 
(1935) 150 — Type: New Britain (Neu Pommern), Gazelle Penins., Ralum, Warburg 20709 (B. 7. 
n.v.). 


ho bo 


Tree 5-18 m. Twigs terete, towards apex 5-10 mm diam., early to late glabrescent 
from light rusty or yellow-brown tomentum with hairs 0.5-1.0 mm long, bark rather 
finely striate, when older not flaking; lenticels present but not distinct. Leaves in 2 
rows, membranous, oblong-lanceolate to lanceolate, broadest towards the base or 
usually + parallel-sided, 30-60 x 7-11 cm, base nearly rounded to 
(short-)attenuate, tip long acute-acuminate, usually 1-2 cm caudate: upper surface 
drying dull olivaceous, minutely, palely punctate-pustulate, lower surface with 
persistent tomentum of pale brown softish dendroid hairs of mixed sizes 0.5-1 mm, 
when shed, not leaving thickened rough hair bases; without brown dots; midrib flat 
above, glabrescent except at the very base; nerves 30-40 pairs, generally + striaght, 
at an angle of 50-70° to the midrib, 8-15(-20) mm apart, thin and flat above; 
beneath, the marginal nerve rather distinct, not very regularly looping; tertiary 
venation lax, indistinct above; petiole 5-16 x 3-4 mm, not or hardly winged; leaf 
bud stout, 4-6 cm long, with dense velvety tomentum with hairs (0.5-) 1 mm. 
Inflorescences woolly pubescent, hairs 1-1.5 mm long, in CO and Q: 2 or 3 times 
ramified, rather many-flowered, 4-15 x 3-10(-12) cm, common peduncle 5-50 mm 
long; bracts broadly ellipsoid, subacute, 5-10 mm long. Flowers in loose clusters of 
3-6 each, perianth 2-valved, glabrous except at the very base, pedicels pubescent 
with hairs 0.3-0.8 mm long, at base inarticulate. Male perianths obovoid-ellipsoid, 
at apex acutish, c. 2-2.3 x 1.5-1.7 mm, pedicel 1(-2) mm long, perianth at anthesis 
split toc. 44, valves c. 0.2 mm thick. Androecium c. 1-1.1 X 1.2-1.3 mm, = laterally 
flattened, broadly rounded above, column at apex narrowly hollowed for c. 4 
(0.2-0.3 mm), anthers c. 10(-12), + completely sessile, erect, mutually touching, c. 
1-1.2 mm long, free apices c. 0.2 mm; androphore slender, 0.3-0.4 mm long. 
Female perianth obovoid, c. 4 x 2.5-3 mm, split at anthesis to c. %4, valves c. 0.3 
mm thick; pedicels 1-1.5 mm; ovary ovoid, c. 2 X 1.5 mm, densely pubescent with 
hairs c. 0.5 mm, style and stigmas minute, + elongate, minutely 2-lobed, c. 0.2 mm 
long. Fruits 2-10 per infructescence, ellipsoid, top obtusish, base broadly rounded, 
2.5-3.0 X 1.5-1.9 cm, pubescent, hairs c. 0.5 mm long, coarse, paler-coloured 
lenticel-like tubercles present, dry valves 3-5 mm thick; stalk 1-3 mm long; perianth 
not persisting. 


Distribution. West and E. New Britain (Gazelle Penins.) 


BISMARCK ARCH. New Britain: LAE 52084; NGF 7060, 7092, 36304 A, 36773, 38152, 44388, 
Warburg 20709 (B, 7 n.v.). 


Ecology. Lowland rain forest, recorded from well-drained pumice terrain, sandy 
soil, ridge forest, at edge of swamp; 0-100 m. Flowering and fruiting apparently not 
seasonal. 


144 Gard. Bull. Sing. 38(1) (1985) 


NOTES 


1. Fieldnotes. A small or medium sized tapered understorey or subcanopy tree; 
bole straight, branches horizontal but drooping terminally; once recorded as slight- 
ly buttressed. Bark dark coloured, mottled, or with short vertical fissures. Wood 
straw, moderately soft. Flowers yellow. Inflorescences (with flowers) recorded as 
erect. Mature fruit green or brown-green. 


2. Related to H. leptantha, H. pulverulenta, and H. hellwigii, the last also 
occurring on New Britain. Distinguished from H. hellwigii by the larger fruits, 
smaller and narrower male flowers (with a somewhat different androecium), and 
the generally more elongated leaves. 


3. Sinclair included the present species in H. hellwigii var. hellwigii. 
4. I have not seen the type, which was apparently lost in B, but Warburg notes 


that the anthers do not reach the top of the androecium and leave a sterile narrow 
appendix. This latter is at variance with the material seen by me. 


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VOL. 38 (Part 2) lst December 1985 


ae . i 
‘ 
io 


re 
3 \ CONTENTS 
we 
OX 
N PAGES 
HOLTTUM, R.E.: 
Two New Species of Tectaria from Limestone in Peninsular Malaysia, with Comments 
UN IN eRe O85 Sn wep crisp te ier cabs ete ce oes vce eet a lity pacsescsvtacverscesees 145-148 
KENG, HSuan: 
eee IshOl occu t lants Of Singapore. (1X) Slee sal i PRC idee cae eee eee deness sees 149-174 


LIM-HO CueEE LEN and LEE SING Kona: 
Micropropagation of Lagerstroemia speciosa (L.) Pers. ............0cccecececeee ee ee ee eneneneeees 175-184 


WILDE, W. J.J.O. bE: 
A New Account of the Genus Horsfieldia (Myristicaceae), Pt3 ............ cece eee eee eee 185-225 


BIDIN, Aziz, and TREVoR WALKER: 
Comparative Anatomy of the Stipe of the Fern Genus Adiantum L. .......................... 227-233 


Published by the Botanic Gardens 

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THE GARDENS’ BULLETIN 


VOL. 38 (Part 2) lst December 1985 
CONTENTS 
PAGES 

HOLTTUM, R.E.: 

Two New Species of Tectaria from Limestone in Peninsular Malaysia, with Comments 

a 8 IE a os a ie Lie clot oA Ry Si Sele 145-148 
KENG, Hsuan: 

Pease Laer See F tants Of Singapore (IX) oe cae oso ce- eon seen ecw ewer reece eee egees 149-174 
LIM-HO CueE LEN and LEE SING Kona: 

Micropropagation of Lagerstroemia speciosa (L.) Pers. .............6..00 cc ccec ence eee e ences eee 175-184 
WILDE, W.J.J.O. DE: 

A New Account of the Genus Horsfieldia (Myristicaceae), Pt3 ........... 0... c cece eee 185-225 
BIDIN, Aziz, and TREVOR WALKER: 

Comparative Anatomy of the Stipe of the Fern Genus Adiantum L. .......................... 227-233 


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Two New Species of Tectaria from Limestone in Peninsular Malaysia, 
with Comments on Some Other Species 


R. E. Holttum 


Royal Botanic Gardens 
Kew, Surrey, England 


EFFECTIVE PUBLICATION DATE: 15 FEB. 1986 


SUMMARY 


Tectaria curtisui and T. translucens are described as new species, with comments on T. coadunata (J. 
Sm.) C. Chr.. T. brachiata (Zoll. & Mor.) Morton and TJ. variolosa (Hook.) C. Chr. Evidence is 
presented that the type specimen of Pieridrys acutissima Ching was probably collected on limestone and 
that the single collection of Hypodematium on limestone in Kelantan represents the species Hypodema- 
ttum glabrius (Copel.) Holttum, comb. nov. 


THE NEW SPECIES 


Tectaria curtisii Holttum sp. nov. 


Stipes usque 70 cm longus, basin versus castaneus, paleis angustis fuscis vestitus, sursum pallidior, 
minute pilosus; lamina usque 60 cm longa, tenuis, pinnis stipitulatis 3-paribus etiam sessilibus vel adnatis 
3-paribus constituta: pinnae infimae ultra 30 cm longae, longe stipitulatae, pinnulas liberas unijugatas 
ferens, pinnula infima basiscopica 20 x 7 cm, profunde lobata, lobis acuminatis lobulatis, lobo infimo 
libero; pinnae suprabasales 30 cm longae, pinnulis liberis unijugatis subaequalibus praeditae: venae in 
areolis angustis costalibus et costularibus ordinatae, venulis retrorsis inclusis. areolis alliis ut in T. 
coadunata sed paucioribus et nonnullis venulis brevibus liberis instructis; costae, costulae venaeque 
subtus pilis patentibus tenuibus 0.2-0.3 mm longis vestitae, pagina inter venas pilis erectis multis 
brevioribus praedita; costae supra pilis brevibus dense vestitae, pili ad et inter venas rares, margines 
(sinubus inclusis) pilis destitutae: sori plerique ad venas breves in areolis terminales; indusia tenuia, 
glabra. 


Typus: near Ipoh, on limestone, C. Curtis 3376, December 1895 (holotypus K: 
isotypus SING). No other specimen is known. 


This species differs from the small Peninsular plants of Tectaria coadunata (J. 
Sm.) C. Chr. in having much larger and more amply branched fronds with abun- 
dant hairs between veins on the lower surface and practically glabrous on the upper 
surface (hairs are lacking even at the sinuses). The new species differs also in the 
presence of small free veinlets arising from the outer veins of the costal areoles and 
directed towards the costae; such veinlets are absent in 7. coadunata. Short free 
unbranched veinlets are also present in other areoles, often directed towards the 
costa; branched included veinlets have not been noticed. 


Tectaria translucens Holttum sp. nov. 


Caudex brevis, erectus; stipes usque 35 cm longus, modice castaneus, sparsim brevipilosus, basi paleis 
usque 15 x | mm, apice capillaceis, vestitus; lamina usque 40 cm longa, tenuis, translucens, pinnis 
stipulatis 2-jugatis etiam sessilibus vel adnatis 2-jugatis constituta: pinnae infimae usque 19 cm longae, 
pinnulis unijugatis praeditae, pinnula infima basiscopica 12 cm longa, basin versus profunde lobata: 
venae in areolis angustis costalibus et costularibus venulis liberis retroversis inclusis ordinatae, venulae 
liberae etiam in areolis alliis pleraeque costam versus currentes, raro furcatae, adsunt: axes frondis 
subtus sparsim brevipilosi, pagina inter venas utrinque glabra: sori plerique ad venas exteriores costular- 
um, in lobis pinnarum distaliter ad venas liberas terminales siti; indusia tenuia, pilis brevibus interdum 
praedita. 


146 Gard. Bull. Sing. 38(2) (1985) 


Typus: Pahang, Taman Negara, on limestone, B.S. Parris and P.J. Edwards 
10450 (K). 


Young plants may have some hairs between veins on both surfaces. 


A distinctive feature of these two new species is the presence of free veinlets in 
the costal areoles. Such veinlets occur also in T. cherasica (Holttum 1981, p. 141) 
and in 7. brachiata (discussed below). This character distinguishes all four species 
from T. coadunata, which has the basic vein-pattern on which Presl (1836) based his 
genus Sagenia. Another difference 1s that in T. coadunata free veinlets in non-costal 
areoles are few, unbranched and outwardly directed, whereas in the other species 
many free veinlets are directed towards the costa and in some cases are branched. 
Branched veinlets are frequent in the sterile fronds of T. brachiata but rare in T. 
cherasica and T. translucens and they have not been noted in T. curtisii. Apart from 
these species there is, within Tectaria, a rather clear distinction between the species 
which show the Sagenia pattern and those which have abundant branched free 
veinlets in areoles (see Holttum 1983, p. 108 and fig. 1). It is notable that in SE 
Asia there is a species (7. fuscipes (Bedd.) C. Chr.) which has Sagenia-type 
venation in sterile fronds but free veins in fertile ones. This bridges the gap between 
species with anastomosing veins and those with free veins which Ching and others 
have included in a genus Ctenitopsis. A majority of such free-veined species occur 
in mainland SE Asia and the Philippines. Combining this information with the fact 
that there are far more species of Tectaria in the Palaeotropics than in the Amer- 
icas, I suggest that the genus Tectaria originated in SE Asia. Taxonomically, it is 
unfortunate that the type species of the genus is a West Indian one. 


TECTARIA VARIOLOSA AND T., BRACHIATA 


In Holttum 1981, p. 137, the name 7. variolosa is placed as a synonym of T. 
brachiata, but in fact the type specimens of the two differ in venation, and the 
characters specified for 7. brachiata in the key on pp. 133-134 are those of T. 
variolosa. 


In both species the pinnae of fertile fronds are contracted as compared with 
sterile ones (intermediate fronds may sometimes occur), and in both the only hairs 
between veins on the upper surface are small ones near the sinuses between 
pinnalobes. The pinnae of sterile fronds of T. brachiata are much less deeply lobed 
than those of 7. variolosa and there are many free and branched veinlets both in 
costal and some other areoles (Fig. 1). 


LS, 
i 


Fig. 1. Venation in a lobe of a middle pinna of a sterile frond of A: Tectaria variolosa (Toppin 4183, 
Upper Burma) and B: Tectaria brachiata (Curtis 1608, Penang), both x 2. 


Two new species of limestone Tectaria 147 


The name 7. variolosa (originally in the genus Aspidium) was copied from 
Wallich’s catalogue and based on Wallich 379. Under this number Wallich included 
specimens from NE India (of which Ching selected one as lectotype) and also from 
Penang. The Penang specimens (of which two sheets are at Kew) are 7. brachiata. 
Accepting Ching’s choice of a type, T. variolosa occurs from NE India southwards 
into Burma, Thailand and Vietnam. Specimens so named from Hainan and Taiwan 
seen by me have broader fertile pinnae with a more ample venation and a greater 
number of smaller sori; they are more like 7. subtriphylla (Hook. & Arn.) Copel. 
and need further study. It is possible that the type of Phlebigonium impressum Fée 
(Griffith, Pl. Indic. 34, now at Rio de Janeiro; see Windisch 1982, p. 59) is 
conspecific with the type of 7. variolosa; if this should be established, Fée’s name 
(1852) is the older and his specific epithet should be substituted for variolosa. 


T. brachiata is widely distributed, but specimens are known from few localities. 
There are several collections from Peninsular Thailand and the northern part of 
Peninsular Malaysia; those for which a habitat is recorded indicate granite rocks, 
not limestone. The other specimens known to me are from Java (including the 
type), the Tenimbar Islands (Buwalda 4262) and the Cape York Peninsula in 
Queensland (Brass 19445, Coveney 7146). The species is evidently adapted to a 
continuously warm climate with a regular dry season. 


Key to the species of Tectaria mentioned in this paper 


1. Free included veinlets lacking in costal areoles 
2.eipper surface Dearine many hairs DEtWEEN VEINS. 2:1... coz ese es nee dine op eceennes sees nee T. coadunata 
2. Upper surface lacking hairs between veins except a few near sinuses between pinna-lobes ........... 
Als wept ap A aes nad «Sm Bay 23 ead bake Abas Cred, Naeeiane aie rent od ws Bnei aE ro a A ne T. variolosa 

1. Free included veinlets present in costal areoles 
3. Fronds dimorphous; free veins present in costal and other areoles in sterile fronds, forked ones 


YOO SSE ee AS UC) ts ee ern eee oe eee ee er oe T. brachiata 
3. Fronds not or little dimorphous; fewer free veins in areoles, forked ones infrequent; limestone 

plants 

4. Scales at base of stipes thin, light brown, becoming crumpled ......................265. T. cherasica 


4. Scales at base of stipes firm, dark, much narrowed towards their tips 
5. Fronds to 60 cm long; a free tertiary leaflet present on basal pinnae; lower surface copiously 


Ce EPI SETS Zin (>) a ea aoe ae on eon ae ae ere 2 eee or T. curtisii 
5. Fronds smaller, no free tertiary leaflet; lower surface between veins glabrous in fronds of 
i ey Ag be Sipe De be a an peel Aelia ie 2 toe gine ile SR Ce one T. translucens 


PTERIDRYS ACUTISSIMA CHING 


This species is described in Holttum 1955, p. 531. The type was collected by 
Mohamed Haniff on a journey to Gunung Korbu with B.H.F. Barnard (Forestry 
Dept.) in 1909. The route was by way of the Korbu River, G. Yong Blar and G. 
Bal; thus he passed near limestone hills. The labels on the specimens at Singapore 
and Kew were written by H.N. Ridley, who named them Lastrea syrmatica and 
wrote on them the altitude 6000 ft, giving them a number (14142) in his own series. 
Haniff’s labels are lost. No other collection of this species has since been reported. 
Recently, however, Dr. B.S. Parris collected an almost identical specimen in 
Gunung Mulu National Park, Sarawak, at an altitude of 100 m, ‘‘on slopes of bat 
guano in cave mouth’’. 


No specimen of this genus has been collected at a high altitude in Malesia, and 
the related species P. syrmatica (Willd.) C. Chr. & Ching, widely distributed, is 
nearly always recorded as growing on limestone. 


148 Gard. Bull. Sing. 38(2) (1985) 


THE GENUS HypPoDEMATIUM 


There is only one record of the existence of this genus in Peninsular Malaysia; the 
specimen is M.R. Henderson 29682, from Gua Teja, Kelantan. In Holttum 1955, p. 
501, this is named H. crenatum (Forsk.) Kuhn, with a note that it differs from the 
typical form of that species which was first described from Arabia and is widely 
distributed in Mainland Asia. On comparison with other specimens in the herbar- 
ium at Kew, I find that the Kelantan specimen agrees closely with the type of 
Dryopteris glabrior Copel., collected by C.J. Brooks at Bidi, Sarawak in 1908. 
Copeland’s species was not transferred to Hypodematium by Ching in his paper of 
1935 (probably he did not know of it). The tronds are much more open in branching 
than those of T. crenatum and the leaflets are thin and quite flat when dried. 
Acicular hairs are rather few, and capitate hairs are also present. The new com- 
bination Hypodematium glabrius (Copel.) Holttum is therefore proposed 
(basionym: Dryopteris glabrior Copel., Phlip. Journ. Sci. 5C (1910) 283). 


In Malesia all recorded plants of the genus have been found on limestone, and 
though they may be locally abundant few specimens have been collected. In the 
herbarium at Kew are three other specimens from Sarawak, all H. glabrius; from 
Sumatra are two collections which appear to represent two different species; one 
specimen from New Guinea which has very abundant very long hairs; and a 
specimen from the Philippines which appears to be true H. crenatum, said by 
Copeland to be known from several localities. A new study of the genus in Malesia 
is desirable, but more field work would first be needed. 


The genus is isolated taxonomically and its affinity is uncertain. Chromosome 
counts show the base numbers 40 and 41. Ching at first suggested an affinity with 
Cystopteris, but later thought Lastreopsis more likely. The frond-form resembles 
that of Lastreopsis but hairs and scales are very different. Recently Iwatsuki (1964) 
suggested an alliance to Athyrioid ferns, perhaps nearest (but not near) Woodsia. 


LITERATURE CITED 


Ching, R.C. (1935). On the genus Hypodematium Kunze. Sunyatsenia 3: 3-15. 

Holttum, R.E. (1955). A Revised Flora of Malaya, 2, Ferns of Malaya. Singapore, 

~ Government Printer. 

______ (1981). The genus Tectaria Cav. in Malaya. Gard. Bull. Singapore 34: 
132-147. 

(1983). The fern-genera Tectaria, Heterogonium and Ctenitis in the Mas- 
carene Islands. Kew Bull. 38: 107-130. 

Iwatsuki, K. (1964). On Hypodematium Kunze. Acta Phytotax. Geobot. 21: 43-54. 

Presl, K.B. (1836). Tentamen Pteridographiae. Prague. 

Windisch, P. (1982). Specimens from Feé’s pteridological collection at the Botanic 
Gardens of Rio de Janeiro. Amer. Fern Journ. 72: 56-60. 


Annotated List of Seed Plants of Singapore (IX)* 


HsuAN KENG 
Department of Botany, National University of Singapore 


Index to Families 


Page Page 
CRIB AMUEAS ein ons <fe0525~-<~ 166 DrRRRGCMMCCAG. 8 cccc--. ces oe .. 167 
(opemoncee 952 p654<-5--,2--) 166 US DTS 2 ee ee ea 149 
ee ee eee eae 167 iyMGIAGedes ce. 4 2-5-2282 5-- 167 


II. Angiospermae-Dicotyledons (cont’d) 


125. RUBIACEAE 


Synoptic key to the generat 


1. Ovary containing numerous (rarely few) ovules in each locule 
2. Fruit dry, capsular 


3. Flower-clusters in globular heads ..........................04. Adina, Nauclea, Neonauclea, Uncaria 
3. Flowers in cyme, corymbose or paniculate 
Pees On dees OF Wondy Cumbre (1.55.50 2 227-2. 6 eack Sncn cess o< 02 Coptosapelta, Mussaendopsis 
4. Herbs or herbaceous twiner!s .................--. Argostemma, Dentella, Hedyotis, Ophiorrhiza 


2. Fruit drupaceous or baccate 
5. Corolla-lobes valvate in bud 


om OE UMP 2 8 yo" 2 eta dpe aed elt atte I ae eee an Lecananthus, Lucinaea 

6. Flowers in cymose clusters, corymbose or paniculate ..... Mussaenda, Mycetia, Urophyllum 

5. Corolla-lobes twisted in bud ............... Diplospora, Gardenia, Hypobathrum, Jackia, Randia 
Scyphiphora 

1. Ovary containing a single (rarely 2 or few) ovule in each locule 
7. Epiphyte with tuberous stems tenanted by ants .........................0.. Hydnophytum, Myrmecodia 
7. Not as above 
8. Corolla-lobes imbricate or twisted in bud .............. Ixora, Coffea (introduced), Gardeniopsis, 


Guettarda, Pavetta, Tarrena 
8. Corolla-lobes valvate in bud 


fe eae ee ee eee Prismatomeris, Timonius 
9. Flowers bisexual 
10. Flowers 1-3 together on a terminal, slender stalk; creeping herbs ................ Geophila 


10. Flowers many in compact umbels or heads, terminal and axillary; trees, shrubs or 


SOO CUNIIIOTS 55.5.5. - osu s9- ores canes Cephaelis, Coelospermum, Gynochthodes, Morinda 
10. Not as above 


11. Flower cymes in axillary fascicles 


pe LS Bb Ls SEA aie 6 0 Rte a Canthium, Lasianthus, Saprosma 
Sia en ACOCOMUS JO SIR pitt dito: 08, ot iil yee A. Borreria, Diodia 


11. Flower-cymes in corymbs or panicles ................. Chassalia, Paederia, Psychotria 


* Continued from Gdns’ Bull. Sing. 36: 124, 1983. . . 
The author is indebted to Dr. Richard T. Corlett and Mr. Hugh T.W. Tan for going through the entire 
manuscript and the Rubiaceae respectively, and for their many suggestions. 


+ Based on Ridley, Fl. Mal. Pen. 2 (1923) 3, modified and simplified; some cultivated genera are not 
included. 


149 


150 Gard. Bull. Sing. 38(2) (1985) 


Adina rubescens Hemsl. (alt. name: Pertusadina eurhyncha (Miq.) Ridsdale) 
Tall tree, the trunk full of elliptic holes so the bark appears reticulate; leaves 
elliptic, 6-10 cm long; flowers in small heads, 6-8 mm across, usually 3 heads 
together on a slender peduncle. Scattered in forests; Bukit Timah (Keng & 
Jumali 3483). Vern. Berambong. 


Borreria alata (Aubl.) DC. (= B. latifolia K. Sch.) 
Fleshy herb, to 60 cm tall; stems winged; leaves obovate or elliptic, 2-7 cm long, 
hairy on both sides; flowers in axillary clusters; corolla 3-6 mm long, white; 
fruit subglobose, of two 1-seeded cocci, hairy. A weed of West Indian and S. 
American origin. 


Bor. articularis (L.f.) F.N. Will. (= B. hispida K. Sch.) 
Diffused herb, about 30 cm high; hairy; leaves obovate or oblong, 1.5-4 cm long; 
flowers in axillary clusters; corolla 4.5-7 mm long, pale purple. In waste ground 
and sandy spots; Changi (Md Nur 29739). 


Bor. laevicaulis (Miq.) Rid. 
Slender herb, to 45 cm long, glabrous, often tinted purple; leaves sessile; oblong- 
lanceolate, 1.5-3 cm long; flowers in dense axillary clusters; corolla 2-3 mm long, 
white. In roadsides and sandy waste places; Pulau Ubin (Furtado 18343). 


Bor. setidens (Miq.) Bold. 
Diffused herb, sometimes creeping, 5-20 cm tall; branches winged; leaves lan- 
ceolate or ovate, 0.8-1.5 cm long; flowers in small axillary clusters; corolla very 
short, 2-3 mm long, white. In waste ground, Pasir Panjang (Ridley 8109). 


Canthium confertum Korth. 
Small tree, to 10 m tall, glabrous; leaves leathery, elliptic, 5-15 cm long, nerves 
3-4 pairs; flowers in small clusters, 1.5-2 cm across; corolla 2-3 mm long, 
short-tubed, 5-lobed. Along tidal rivers near the sea, P. Tekong (Ridley 4893). 


Canth. dicoccum (Gaertn.) T. & B. (= C. didymum Gaertn.) 
Small tree, 7-8 m tall; leaves thin-leathery, ovate-lanceolate or elliptic, 6-10 cm 
long, nerves 4-5 pairs; flowers in clusters, 3-5 cm across. In forests or in man- 
groves, Pulau Serimbun (Sinclair 39530), Chua Chu Kang. 


Canth. glabrum BI. 
Small tree to 13 m tall, glabrous; leaves ovate, 12-18 cm long; flowers in small 
cymose clusters; corolla 3-4 mm long, 5-lobed; berry ellipsoid, 2-3 cm long, 
greenish, with 2 flattened stones inside. In forests; Bukit Mandai (Ridley 4434), 
Tuas. 


Canth. horridum BI. 
Spiny shrub, pubescent; spines straight or curved, 3-5 cm long; leaves ovate or 
elliptic, 2-3 cm long; nerves 3-4 pairs; flowers in cymose clusters; corolla 2-3 mm 
long, pale green. In forest edges or open places; Gardens’ Jungle; Bukit Timah 
(Ridley s.n. in 1893). 


Canth. molle K. & G. 
Spiny climber, velvety; spines decurved, 1.5-2 cm long; leaves lanceolate to 
ovate, 5-7 cm long, brown-hairy; nerves 4-5 pairs; cymes small, axillary. In 
forests, Gardens’ Jungle, MacRitchie Reservoir (Sinclair SF 39147). 


Seed plants of Singapore 1X 151 


Cephaelis singaporensis Ridley 
Low shrub; leaves oblong, 2-6 cm long, narrowed at base; nerves 14-16 pairs; 
cymose heads 2-2.5 cm across; corolla trumpet-shaped, 2-3 cm long, yellow; 
drupe light blue, flattened. In forests; Bajau (Ridley 4966). 


Chassalia curviflora (Wall.) Thw. (= C. chartacea Wall.) 

Small shrub, 1-1.5 m tall, glabrous; leaves membranous, variable elliptic to 
oblanceolate, 6-20 cm long, nerves 5-6 pairs; cymes terminal, 3-5 cm long, 
peduncles and its branches purple or white tinted purple; flowers subsessile; 
corolla 1.5-2 cm long, 5-lobed. Berry ellipsoid or globose, 5-6 mm across, 
2-seeded, seated on the swollen white branches of the inflorescence. In forests; 
Bukit Mandai (Ridley s.n. in 1889). (The generic name often spelled as Chasalia; 
some authors have reduced it to Psychotria). 


Chas. pubescens Rid. 
Small shrub, much branched, hairy; leaves lanceolate,.12-15 cm long; pubescent 
beneath; nerves 7-8 pairs; compound cymose dense, 2-3 cm long, hairy; flowers 
in small clusters, rosy white; corolla 6-8 mm long. In forests; once collected at 
Bukit Timah (Ridley s.n. in 1884). 


Coelospermum scandens BI. 
Slender climber; leaves elliptic or obovate, 6-12 cm long; nerves 4-7 pairs; 
flower-cymes umbellate, fragrant, arranged in a small terminal panicle, 3-4 cm 
long; corolla 2-3 mm long, 5-lobed, white. Berry depressed globose, 1-1.5 cm 
across, 2-4 seeded. Climbing on trees in forests; Changi (Ridley 5926), Bukit 
Timah. (The generic name sometimes spelled as Caelospermum). 


Coffea liberica Bull. ex. Hiern. 
Stout herb or small tree; leaves obovate, thick-leathery, 16-35 cm long, nerves 
6-12 pairs; cymes axillary, subsessile; corolla white, 2.5-3 cm long, 6-8 lobed; 
berry subglobose, 1.5-2.5 cm long, red, with 2 flattened stones inside. Native to 
W. Africa, formerly planted. 


Cof. robusta Linden ex De Wild. 
Stout shrub; leaves broadly oblong, 15-30 cm long; nerves 10-12 pairs; cymes 3-5 
flowered, axillary; corolla 1.5-2 cm long, 5-7 lobed; berry ovoid-globose, 1-1.5 
cm long, red. Native to tropical Africa, formerly planted. Some authors have 
reduced this species to a variety of Cof. canephora Pierre ex Froehner. 


Coptosapelta griffithii Hook.f. 
Climbing shrub; leaves ovate-orbiculate, 4-6 cm long; nerves 3-4 pairs, hairy 
beneath; panicles terminal and axillary; corolla salver-shaped, 1.5-2 cm long, 
white, throat woolly, 5-lobed; capsule obovoid, pubescent, 0.8-1 cm long, seeds 
numerous. In forests; Gardens’ Jungle, Chua Chu Kang (Che Mat 6891). 


Copt. parviflora Rid. 
Lofty climber; leaves elliptic, narrowed on both ends, 6-8 cm long; nerves 4-5 
pairs; panicle terminal, spreading, 6-8 cm long; corolla 6-8 mm long, green; 
capsule obovoid, glabrous, 2-4 mm long. Climbing to tops of trees in forests; 
Bukit Timah (no specimens available). 


152 Gard. Bull. Sing. 38(2) (1985) 


Copt. tomentosa (BI.) Val. ex K. Heyne (= C. flavescens Korth.) 
Lofty climber; leaves ovate or elliptic, 5-10 cm long, nerves 4-5 pairs, soft hairy 
beneath; corolla 2-2.5 cm long, white, fragrant; capsule obovoid, 4-6 mm long, 
glabrous. In forests, climbing on trees; Bukit Timah (Ridley 14117). 


Dentella repens (L.) J.R. & G. Forst. 
Creeping branched herb; leaves oblong to elliptic, 4-10 mm long; flowers solit- 
ary, in one axil of the paired leaves, 5-merous; corolla 0.5-1.5 cm long, yellowish; 
fruit ovoid, 2-3 mm long, dry, indehiscent; seeds numerous. In waste ground and 
damp places; Tanglin, Kallang, Sembawang (Keng et al 4070). 


Diplospora malaccensis Hook.f. 
Small tree, to 10 m tall, glabrous; leaves elliptic, 5-15 cm long; flowers 4-5 in 
small axillary clusters; corolla tubular, 2-3 mm long, 4-lobed, greenish white; 
berry globose, orange, few-seeded. In forests; Water Catchment Area, Bukit 
Timah (Maxwell 8146). 


Diodia ocymifolia (Willd. ex R. & S.) Bremek. 
Rough herb, to 1 m tall; leaves oblong-ianceolate, 3-7 cm long; nerves 5-8 pairs, 
hairy on nerves and near margin; flowers 6-many, in axillary clusters; corolla 2-4 
mm long, 4-lobed; fruit of two 1-seeded cocci. A native of S. America; in damp 
places, forest edges and grassland. 


Diodia sarmentosa Swartz 
Herb; leaves oblong-ovate, 3-6 cm long, nerves 3-5 pairs, scabrous above, nerves 
hairy beneath; flowers 1-8, in axillary clusters. In forest edges and grassland. 
Native to S. America. 


Gaertnera grisea Hook.f. ex Clarke 

Shrub, to 1.2 m tall; leaves thin leathery, oblong or oblanceolate, apex acumin- 
ate, 20-35 cm long, densely hairy beneath, nerves 7-8 pairs; petiole 6-8 mm long, 
thick; stipule-sheath tubular, 2-2.5 cm long; cymes terminal and axillary, 6-8 cm 
long, hairy; corolla white, cylindric, 6-8 mm long; lobes valvate. In forests; Bukit 
Timah, Bukit Panjang, Ponggol (Burkill 7613), Changi. This genus differs from 
almost all the rubiaceous genera in having a superior ovary, therefore it was 
classified under Loganiaceae in Ridley’s Flora. 


Gaert. obesa Hook.f. ex Clarke 
Stout, fleshy shrub, 1.2 m tall; leaves fleshy leathery, oblong or lanceolate, apex 
short-acuminate, 25-35 cm long; nerves 9-12 pairs; stipule-sheath 3-4 cm long; 
cymes dense, umbellate, 4-5 cm across; corolla white, 6-8 mm long, sessile or 


subsessile, densely hairy in the mouth. In forests; Gardens’ Jungle, Tuas, Chua 
Chu Kang (Ridley 2680). 


Gaert. vaginans (DC.) Merr. (= G. acuminata Benth.) 
Shrub; leaves membranous, elliptic, apex acute, 7-9 cm long; stipule-sheath 1-1.5 
cm long; panicles terminal, 8-12 cm long; corolla 5-6 mm long, white, lobes 
acute, as long as the tube, mouth hairy. (In forests, collected in Singapore only 
once (Wallich 8342, not seen). 


Seed plants of Singapore 1X 153 


Gaert. viminea Hook.f. ex Clarke 
Slender shrub, 1 m tall; leaves membranous, narrowly lanceolate, apex caudate, 
6-9 cm long, nerves 6-7 pairs; cymes 4-6 cm long, 3-chotomously branched; 
corolla short-cylindric, 5-6 mm long, white, lobes 4, broadly ovate, as long as the 
tube. In forests; Gardens’ Jungle, Bukit Mandai, Changi. 


Gardenia carinata Wall. ex Roxb. 
Small tree to 7 m tall; leaves thin-leathery, obovate or oblanceolate, 10-40 cm 
long; flowers solitary, in upper axils; calyx tube 1.5 cm long, 5-6 ribbed; corolla 
golden yellow, the tube 2.5 cm long, the limb 6-7 cm across, 6-9 lobed; fruit 
ellipsoid, 3-4 cm long, ribbed, crowned with the calyx lobes. Introduced from N. 
Malaya, planted as a roadside tree. 


Gard. griffithii Hook.f. 
Shrub or small tree, to 7 m tall; leaves obovate, thick-leathery, narrowed at base, 
12-20 cm long; nerves 12-13 pairs; flowers solitary, terminal; calyx tubular, 6-8 
cm long, the mouth expanded; corolla cylindric, 8-10 cm long, orange; fruit 
globose, woody, 4-5 cm across, crowned by the calyx-lobes. In edge of forests; 
Bukit Timah, Bukit Mandai (Ridley 6673). 


Gard. jasminoides Ellis (= G. angusta Merr.; G. florida L.) 
Shrub, 1-2 m tall; leaves obovate, thick and shining, 8-10 cm long; flowers 
solitary, axillary; calyx tubular, angled or winged; corolla waxy white, turning 
yellowish, 3-5 cm long, 5-10-lobed, fragrant. Native of S. China, in several 
horticultural forms, some double-flowered, sometimes planted in gardens. Vern. 
Bunga Susu, Bunga China, te--té 


Gard. tubifera Wall. 
Shrub, to 2 m tall; leaves oblanceolate, much narrowed at the base, 8-24 cm long, 
nerves 15-18 pairs; flowers terminal; calyx tubular, 1.5-2 cm long; corolla 12-14 
cm long, creamy white turning orange-yellow, fragrant; fruit globose, 3-5 cm 
across. Along river banks in mud; Chua Chu Kang (Ridley s.n. in 1895). 


Gardeniopsis longifolia Miq. 
Shrub, with few stout branches, 2-3 m tall; leaves oblanceolate, 20-30 cm long; 
flowers sessile, in axillary clusters; corolla cylindric, 2-2.5 cm long, 5-lobed, rosy 
white; fruit ovoid or ellipsoid, with 10 ridges, 2-2.5 cm long, crowned with the 
enlarged, incurved calyx-lobes. In dense forests, Bukit Timah (no specimens 
available). 


Geophila pilosa Pears 
Small creeping herb, hairy; leaves ovate-cordate, 1-2.5 cm long; flowers termin- 
al, solitary or 3 together on a slender stalk; corolla small, white, funnel-shaped, 
4-6 lobed; drupe globose, black, with 2 stones inside. Rare in forests; Bukit 
Timah, (Ridley 9516), Reservoir woods. 


Guettarda speciosa L. 
Small tree, 5-10 m tall; leaves obovate, 10-25 cm long, the base heart-shaped; 
cymes axiillary, 4-11 cm long; corolla trumpet-shaped, 6-11 cm long, limb 6-8- 
lobed, white, fragrant; fruit depressed globose, 2-2.5 cm across, greenish, faintly 
and closely ribbed, with a circular calyx-scar on the top. On sandy and rocky 
shores; Pulau Seletar (Samsuri SA 1214). 


154 Gard. Bull. Sing. 38(2) (1985) 


Gynochthodes coriacea BI. 
Woody climber, glabrous; leaves leathery, ovate-lanceolate, 6-10 cm long, tip 
blunt; nerves 4-6 pairs; flowers small, few in an axillary cluster; corolla 2-4 mm 
long, woolly inside; drupe globose, 1 cm across, white, with 2-4 stones inside. In 
forests, climbing on trees; Changi, Gardens’ Jungle, Mandai (Samsuri 1388). 


Gynoch. sublanceolata Miq. 
Slender woody climber; leaves thin-leathery, elliptic-lanceolate, 5-8 cm long, tip 
acuminate, nerves 5-6 pairs; flowers sessile, few on very short axillary branches; 
corolla white; drupe globose, 4-5 mm across. In open country, often near the sea; 
Changi, Pulau Tekong (Goodenough 2836). 


Hedyotis.auricularia L. 
Perennial, to 50 cm high; stems 4-angled, covered with white hairs; leaves 
lanceolate-oblong, 4-8 cm long; flowers in axillary, sessile, dense cymes; corolla 
salver-shaped, 2-3 mm long, white; capsule globose, hairy, indehiscent, few- 
seeded. In open places and road sides; Tuas, Chua Chu Kang (Ridley 4125). 


Hedy. biflora (L.) Lamk. (= Oldenlandia paniculata L., O. biflora L.) 
Ascending herb, to 20 cm high; leaves oblong or ovate, slightly fleshy, 5-10 mm 
long; flowers 3-10 in peduncled and often branched (terminal and axillary) 
cymes; corolla 2-2.5 mm long, white or purplish; capsule ribbed. In waste places; 
Tanglin (Kassim 518). 


Hedy. capitellata Wall. ex G. Don 
Climbing shrub, glabrous; ledves from branches lanceolate to oblong, 3.5-11 cm 
long; terminal panicles consisting of umbellate heads (1-1.5 across); corolla 
creamy coloured, 4-merous, fragrant, hairy inside. Climbing over bushes and 
hedges; Tanglin (Ridley 15429). 


Hedy. congesta Wall. ex G. Don 
Shrubby herb, to 2 m high; leaves leathery, lanceolate or elliptic, 10-15 cm long; 
flowers in axillary dense cymes, sessile; corolla campanulate, 3-4 mm long; fruit 
fleshy, oblong, white.In forests; Gardens’ Jungle, Bukit Timah (Burkill HMB 
1894). 


Hedy. corymbosa (L.) Lamk. (= Oldenlandia corymbosa L.) 
Erect or ascending herb, to 30 cm high; leaves oblong, acute, 1-3 cm long; 
flowers axillary, solitary or 2-8 in peduncled cymes; corolla white or pale purple, 2 
mm long. In sunny places or rocky areas. Singapore (Furtado s.n. in 1924). 


Hedy. dichotoma Koen. ex Roth (= Oldenlandia dichotoma Hook.f.) 
Slender, diffused herb, 30-40 cm tall; leaves lanceolate or oblong, 1.5-3 cm long; 
cymes axillary, and terminal, in several dichotomous branches; corolla bell- 


shaped, 1.5-2 mm long, the lobes spreading. In dry sandy places near the sea; 
Geylang (Ridley 11512). 


Hedy. diffusa Willd. (= Oldenlandia diffusa (Willd.) Roxb.) 
Prostrate herb, to 40 cm high; leaves sessile, linear lanceolate, 1-2 cm long; 
flowers solitary or in pairs, axillary sessile or nearly so; corolla white or purple; 
capsule depressed globose, 2-3 mm long, usually nodding. In open sandy places; 
Chua Chu Kang (Hullett 333), Water Catchment Areas. {1 {i We 4 4 


Seed plants of Singapore IX 155 


Hedy. herbacea L. 
Annual, to 20 cm high; leaves sessile, linear or narrowly elliptic, 1-2.5 cm long; 
flowers axillary, solitary or 2-4 in peduncled cymes; corolla white, 2 mm long. In 
open places, Singapore (Ridley s.n. in 1892). 


Hedy. pinifolia Wall. ex G. Don. 
Prostrate annual, to 30 cm high; leaves thick, sessile, linear to lanceolate, 1-1.5 
cm long; flowers solitary, or 2-4 in peduncled cymes, axillary; corolla 3 mm long, 
white or purple; capsule with long bristles. In sandy places by the sea; Jurong 
(Ridley 8924). 


Hedy. trinervia (Retz.) R. & S. (= Old. trinervia Retz.) 
Prostrate annual, to 20 cm long; leaves thin, obovate, 0.2-0.5 cm long; flowers 
solitary, sessile or nearly so, axillary; corolla 1.5 mm long, white. In open spaces 
with sandy soil; Geylang, Pulau Sudong (Maxwell 82295). 


Hydnophytum formicarium Jack 
Epiphytic small shrub; fleshy tuber irregularly lobed, 16-20 cm across, glabrous, 
tunnelled and perforated forming an ant’s nest; leaves leathery, elliptic, 4-10 cm 
long, subsessile, nerves 7-11 pairs; flowers 3-5, in axillary sessile cymes; corolla 
salver-shaped, 3-4 mm long, white; drupe narrowly ellipsoid, 4-5 mm long, with 2 
stones inside. In secondary forests, often near the sea; Jurong, Changi (Ridley 
303). 


Hypobathrum coniferum (Ridl.) Bakh. f. (= Petunga conifera Ridl.) 
Slender tree, 10 m tall; branches 4-angled, glabrous; leaves leathery, 15-22 cm 
long, nerves 7-9 pairs; flowers small, in pairs on cone-like dense axillary spikes, 
4-angled, 1-1.5 cm long; corolla funnel-shaped, 2-3 mm long, 4-lobed, white. 
Only one tree formerly found in the Gardens’ Jungle (Ridley 10722, type). 


Ixora chinensis Lamk. 
Shrub, branched, to 2 m tall; leaves short-stalked, obovate-oblong, leathery, 
6-10 cm long; flowers in dense cymose corymbs, 5-10 cm across; corolla 3-3.5 cm 
long, 4-lobed, the lobes rounded, yellow turning red. Native to S. China and 
Thailand, cultivated. {j\\ ?} fé 


Ixora coccinea L. 
Branched shrub; leaves sessile, ovate to obovate, 3.5-10 cm long, the base 
heart-shaped, clasping the stem; corolla 3-4.5 cm long, red or sometimes yellow 
or pink, often fragrant; the lobes pointed. Native of India, cultivated. 


Ixora congesta Roxb. 
Shrub or small tree, to 7 m tall; leaves elliptic-oblong, 12-30 cm long; corymbi- 
form inflorescence 15-20 cm across; corolla yellow turning orange or pink. In 
forests, Bukit Timah, Chua Chu Kang. 


Ixora finlaysoniana Wall. ex G. Don 
Shrub or small tree; leaves oblong or oblanceolate, leathery, 10-18 cm long; 
flowers white, fragrant. Native to India and Andaman Islands, sometimes culti- 
vated. 


156 Gard. Bull. Sing. 38(2) (1985) 


Ixora grandifolia Z. & M. 
Bushy tree, to 2 m tall; leaves variable, lanceolate or obovate, 12-20 cm long, 
narrowed at base; corymbiform inflorescence 10-20 cm across; corolla 1-1.5 cm 
long, white tinged with pink; fruit ellipsoid, red, 6-8 mm long. In forests, 
formerly collected from Chua Chu Kang (Ridley 4/20). 


lxora javanica (Bl.) DC. 
Shrub, variable; leaves elliptic oblong, 10-25 cm long, shortly stalked; corolla 


2.5-3.5 cm long, yellow turning red; the lobes bluntly pointed. Native to Java, 
cultivated. 


Ixora lobbii K. & G. 
Shrub, 2-3 m tall; leaves lanceolate or oblong, 10-25 cm long, many (15-25)- 
nerved; corymbiform inflorescence 12-15 cm across; corolla 3-4 cm long, yellow- 


orange turning red, the lobes acute. In forests; Sungei Morai, Gardens’ Jungle, 
Changi (Hullett s.n. in 1893). 


Ixora pendula Jack 
Shrub, to 8 m tall, glabrous; leaves variable, oblong or elliptic, 10-25 cm long; 
corymbiform inflorescence 10-20 cm across; corolla 2-3.5 cm long, rose-red. In 
forests; Changi (Ridley 2868), Gardens’ Jungle. 


Jackia ornata Wall. 
Slender tree, to 10 m tall; leaves leathery, oblanceolate, 15-40 cm long, pubes- 
cent, nerves 10-12 pairs; withering red; stipule-sheath with many long bristles or 
teeth round the edge; flowers small in axillary hanging panicles, 15-35 cm long; 
corolla funnel-shaped, less than 1 cm long, yellowish white; fruit obconic, nut- 
like, 1.5 cm long, crowned by 3 enlarged calyx-lobes. In swampy forests, Kranji, 
Dalvey Road (Ridley 4114). 


Lasianthus appressus Hook.f. 
Small shrub, densely covered with soft yellow hairs; leaves leathery, lanceolate 
or oblong, 5-8 cm long; nerves 6 to 7 pairs; stipules prominent; flowers axillary, 
few together, sessile, surrounded by linear hairy bracts; corolla trumpet-shaped, 
4-5 lobed, 5-6 mm long; drupe globose, 2-3 mm across, blue, hairy, with 2-6 
stones inside. In forests; Chua Chu Kang (Ridley 4122). 


Las. attenuatus Jack (Syn. L. densifolius Miq.) 
Shrub, covered with yellow hairs; leaves membranous, lanceolate-oblong, 5-8 cm 
long nerves, nerves 7-8 pairs; flowers 1-3 in axillary cymes. In forests; Bukit 
Timah, Chua Chu Kang (Ridley 16704). 


Las. chryseus Rid. 
Small shrub, | m tall, covered with appressed yellow hairs; leaves thin leathery, 
lanceolate, 10-12 cm long, nerves 8-9 pairs; cymes 5-6 flowered, sessile. In 
forests: Gardens’ Jungle, Chua Chu Kang, Bukit Timah (Ridley 8126). 


Las. constricta Wight 
Shrub; leaves thin-leathery, elliptic-oblong, 8-11 cm long, nerves 3-5 pairs; 
axillary cymes 4-6 flowered, sessile. In forests; Changi (Ridley s.n. 1892), Kranji, 
Sungei Buloh. 


Seed plants of Singapore LX 157 


Las. cyanocarpus Jack 
Shrub, hairy; leaves, leathery, oblanceolate to oblong, 10-16 cm long, the base 
narrowed, unequal, nerves 7-10 pairs; axillary cymes 3-4-flowered, sessile. In dry 
sandy places near the sea; Changi, Pulau Ubin (Ridley 9499). 


Las. ellipticus Wight 
Shrub, hairy; leaves thin-leathery, ovate or elliptic, 10-16 cm long, nerves 6-8 
pairs; cymes few-flowered. In forests, Chua Chu Kang (Ridley 6146), Pasir 
Panjang. 


Las. griffithii Wight 
Small shrub, sparsely pubescent; leaves leathery, oblanceolate or elliptic, 20-30 
cm long, nerves 13-15 pairs; flower-clusters 2-2.5 cm across; corolla white. In 
forests; Chua Chu Kang (Ridley 4/21). 


Las. maingayi Hook.f. 
Small shrub, young parts covered with yellow hairs; leaves membranous, narrow- 
ly elliptic, 12-17 cm long, nerves 4-5 pairs; cymes few-flowered. In forests; Bukit 
Timah (Ridley 12550), Gardens’ Jungle. 


Las. ridleyi K. & G. 
Shrub, soft-pubescent; leaves thin-leathery, oblong or elliptic, 12-20 cm long: 


flower-clusters 1-1.5 cm across. In forests, Gardens’ Jungle (Ridley 4894), Bukit 
Timah. 


Las. scabridus K. & G. 
Shrub, densely hairy; leaves leathery, elliptic-oblong, 10-20 cm long, nerves 


10-12 pairs; flower-clusters dense globose, 2-1.5 cm across. In forests; Jurong 
(Burkill 713). 


Las. singaporensis K. & G. 
Shrub, soft-hairy; leaves thin-leathery, elliptic, narrowed to both ends, 12-15 cm 
long; cymes 4-6 flowered. In forests; Bukit Timah (Ridley s.n. in 1891), Seletar. 


Las. stipularis Bl. 
Slender shrub, glabrous; leaves thin-membranous, lanceolate or oblong. base 
narrowed, decurrent to the petiole, 12-16 cm long; stipules broad-deltoid, com- 
pletely covering the flowering heads. In forests; Bukit Timah, Bukit Panjang 
(Ridley 12547). 


Las. tomentosus BI. 
Shrub, densely hairy; leaves leathery, oblong-lanceolate, 5-8 cm long, nerves 
7-10 pairs; flowers in dense axillary cymes, sessile. In forests: Woodlands, Kranji 
(Ridley s.n. in 1891). 


Lecananthus erubescens Jack 
Woody climber; branches 4-angled, glabrous; leaves fleshy membranous, nar- 
rowly oblong or elliptic, 7-20 cm long; flowers in axillary involucres, round or 
oblong, clusters, 2-2.5 cm across; corolla funnel-shaped, white, tinted purple, 
5-lobed. In wet forests, creeping on trees; Jurong, Kranji, Chua Chu Kang 
(Ridley 3814). 


158 Gard. Bull. Sing. 38(2) (1985) 


Lucinaea membranacea King 
Epiphytic climber, pubescent; leaves membranous, oblong or elliptic, 8-12 cm 
long, nerves 8-9 pairs, midrib red beneath; flowers sessile, 8-10 in terminal and 
axillary heads, 1.5-2 cm across; heads 1-3 together, peduncled; corolla funnel- 
shaped, 2-3 mm long, 5-6 lobed; berry subglobose, fused at base forming a small 
head 6-8 cm across, red. In forests; Bukit Mandai (Ridley s.n. in 1891), Chua 
Chu Kang. 


Luc. morinda DC. 
Scandent shrub sometimes epiphytic, glabrous; leaves leathery, elliptic- 
lanceolate to ovate, 3-6 cm long, nerves 8-11 pairs; flower-heads 2 cm across, 
peduncled, 1-6 heads together terminal; corolla 5-6 mm long, white, fragrant; 
fruiting heads 1.5-2 cm across. In open sandy spots near the sea; formerly 
collected by W. Jack at Thomson Road, no specimens available. 


Morinda citrifolia L. 
Shrub or small tree, 3-4 m tall; leaves membranous, ellliptic, 15-20 cm long; 
flowers sessile, in globose heads, 1.5-2 cm across, terminal and axillary; corolla 
cylindric 1.5 cm long, white; fruit-cluster oblong-ovoid, 5-7 cm long, whitish, 
formed by the fusion of small fruits with their succulent calyces and the axis. 
Leaves are edible and fruits are used in local medicine; according to Ridley, it 
was probably introduced from Moluccas. Vern. Mengkudu. 


Mor. ridleyi (K. & G.) Ridl. 
Large climber; leaves obovate or oblong, base round, 6-8 cm long, densely 
red-hairy beneath, nerves 4 pairs; small heads 5-6 mm across, umbellate; berry 
black. In forests; Gardens’ Jungle (Ridley 6470). 


Mor. rigida Miq. 
Stout climber, soft-hairy; leaves thick-leathery, elliptic, 6-10 cm long, nerves 
18-20 pairs; fruiting heads 2-2.5 cm long, green. Usually near the sea; Changi, 
Kranji (Ridley 4126). 


Mor. umbellata L. 
Climbing shrub; leaves thin-leathery, lanceolate or elliptic, 7-12 cm long, glab- 
rous, nerves 5-7 pairs; flower-heads subglobose, 5-6 mm across, umbellate; 
fruiting heads orange. In open sandy places; Kranji (Ridley s.n. in 1892), Changi, 
Pulau Ubin. 


Mussaenda erythrophylla Schum. & Thonn. 
Shrub, drooping or climbing, 2-4 m high; the calyx-lobes directed to the outside 
of inflorescence, enlarged, petaloid, obliquely ovate, 5-10 cm long, bright red 
above, pale red beneath. Native to tropical Africa, cultivated in gardens. 


Mus. glabra Vahl 
Climbing shrub, glabrous; leaves leathery, variable, lanceolate or elliptic, 6-9 cm 
long, nerves 5-6 pairs; cymes dense, terminal; calyx bell-shaped, the enlarged, 
petaloid lobe broadly ovate, 8-10 cm long and wide, white; corolla trumpet- 
shaped, dark red or orange-red. In secondary forests; Chua Chu Kang (Hullett 


846). BER TE 


Mus. flava (Verdcourt) Bakh. f. (= M. luteola Delile) 
The enlarged, petaloid lobe lemon yellow, 2.5-3.5 cm long. Native to tropical 
Africa. 


Seed plants of Singapore IX 159 


Mus. mutabilis Hook.f. 
Glabrous shrub; leaves membranous, elliptic to ovate, 12-18 cm long, nerves 8-9 
pairs; compound cymes terminal; calyx cylindric, the 5 lobes all narrow lan- 
ceolate, similar, not enlarged; corolla tubular, 5-lobed, orange or bright red, 
fading orange-yellow. In forests; Seletar (Goodenough 1643). 


Mus. philippica L. C. Rich. 
Shrub or rarely small tree, to 5 m tall; branches drooping; flowers prolifically 
most of the year. Several hybrid forms: only one of the calyx-lobes enlarged, pale 
pinkish (“‘Alicia’’), or all five of the calyx-lobes enlarged, creamy white (‘‘Dona 
Aurora”) or pink (“Dona Luz’’). Introduced from the Philippines, widely 
planted. 


Mussaendopsis beccariana Baill. 
Large tree, glabrous; leaves leathery, nearly orbicular, 8-15 cm long, nerves 5-6 
pairs; cymes in panicles, 15-20 cm across; calyx bell-shaped, usually with 5 small 
lobes and 1 large, the large lobes obovate, white, 2-3 cm long; capsule oblong, 
1-1.5 cm long. In forests, Jurong, Sembawang, Chua Chu Kang (Goodenough 
1850). 


Mycetia malayana (Wall.) Craib (= Adenosacme malayana Wall.) 
Shrublets, 50-70 cm high; bark white, shiny; leaves membranous, elliptic- 
lanceolate, 15-25 cm long, densely hairy; flowers in terminal cymose panicles, 
5-15 cm across; corolla yellow, very short (2-3 mm long), 5-lobed; berry subglo- 
bose, 2-3 mm across, white, pulpy. In dense forests; Bukit Timah (Holttum 
19794). 


Myrmecodia armata DC. (= M. tuberosa Bl.) 
Epiphytic shrub, tuber 10-20 cm across, strongly spiny, tunnelled and perforated 
forming an ant’s nest; leaves leathery, elliptic, base narrow, 7-12 cm long, nerves 
7-10 pairs; flowers solitary or few together at base of cup-shaped cavities in stem; 
sessile; corolla cylindric, 3-4 mm long, 4-lobed; drupe oblong, orange, 2 cm long, 
4-8 seeded. On trees near the sea, not common; Jurong (Ridley 3554), Bukit 
Timah. 


Nauclea officinalis (Pitard) Merr. & Chun (= Sarcocephalus junghuhnii Miq.) 
Bushy tree, to 10 m tall; leaves leathery, elliptic to obovate, shortly acuminate. 
10-15 cm long, nerves 5-7 pairs; flowers sessile, joined by the fused calyx-tubes 
into axillary globular heads, 1-1.5 cm across; peduncles stout, 3-7 cm long; 
corolla funnel-shaped, 2-3 mm long, creamy yellow, fruiting heads globose, 1.5-2 
cm across. In forests; Upper Thomson Road (Sinclair SF 40379). 


Nauc. subdita (Korth.) Steud. 
Small trees, 4-10 m tall; branches glabrous; leaves thin-leathery, elliptic to 
broadly oblong, 8-14 cm long, nerves 5-7 pairs; flowers in peduncled heads, 1.5-2 
cm across; corolla 5-6 mm long, pale yellow. In forests; Bukit Timah (No 
specimens available). 


Ophiorrhiza singaporensis Rid. 
Herb, 10-30 cm long; stems succulent, hairy; leaves lanceolate-oblong, 10-15 cm 
long; flowers on the upper side of the branched terminal inflorescence; corolla 
cylindric, white, 4-5 mm long, 5-lobed; capsule strongly flattened, broadly heart- 
shaped, 4-5 mm long, hairy. In forests; Bukit Timah, Chua Chu Kang (Ridley 
s.n. in 1890). 


160 Gard. Bull. Sing. 38(2) (1985) 


Paederia scandens (Lour.) Merr. (= P. foetida L.) 
Twining slender shrub, foetid; leaves opposite, lanceolate or ovate, 5-8 cm long, 
nerves 4-5 pairs; flowers in terminal or axillary cymose panicles, 10-15 cm long; 
corolla funnel-shaped, pubescent, 1-1.5 cm long, violet; fruit orbicular, thin- 
walled, 4-6 mm wide, orange. In open places. 


Paed. verticillata BI. 
Like the above species, but stems stouter and the leaves thicker and larger (6-10 


cm long), opposite or in whorls of three; fruit white, flattened. In forest edges; 
Chua Chu Kang (Ridley 3647). 


Pavetta indica L. 
Shrub or small tree, Ixora-like; leaves oblanceolate or elliptic, 7-15 cm long, 
velvety; flowers in corymbose cymes, 5-10 cm across; corolla 1.5-2 cm long, 
4-lobed, white, style projecting 1.5-2.5 cm beyond the corolla-tube. In forests; 
Chua Chu Kang, Kranji (Cantley 2699). A highly variable species, this plant 
sometimes considered to be a variety of P. indica, namely P. indica L. var 
canescens (Wall.) Ridl. 


Pentas lanceolata (Forsk.) Deflers. (= P. carnea Beth.) 
Herb or subshrub, hairy; leaves opposite, ovate, pointed, 3-8 cm long; cymes 
congested in terminal corymbose or head-like, 6-8 cm across; corolla tubular, 2 
cm long, 5-lobed, purple, pink or white. Native to tropical Africa, sometimes 
cultivated in pots. 


Prismatomeris tetrandra K. Schum. (= P. malayana Ridl.) 
Shrub, 2-3 m tall, glabrous; leaves thin-leathery, variable, lanceolate to elliptic, 
6-12 cm long, nerves 5-7 pairs; flowers with slender pedicels (1-1.5 cm long), in 
terminal sessile fascicle; corolla slender, white, fragrant, 1.5 cm long, 5-lobed; 
drupe globose, 1- or 2-seeded. In open places: Changi (Ridley s.n.in 1891), Chua 
Chu Kang. 


Psychotria angulata Korth. 
Erect shrub, glabrous; leaves leathery, oblanceolate, 9-15 cm long, nerves 8-10 
pairs; cymes paniculate, terminal, 2-3 cm across; corolla 2-3 mm long, tubular, 
5-lobed, the lobes silky inside; drupe red or black, ellipsoid, 4-5 mm long, 
2-seeded, the seed grooved. In forests; Changi (No specimens available). 


Psych. cantleyi Rid. 
Slender climber; leaves lanceolate or ovate, 5-8 cm long, nerves 5-6 pairs; flowers 
in compact cymes. In forests; Chua Chu Kang (Ridley s.n. in 1892). 


Psych. griffithii Hook.f. 
Low shrub, glabrous; leaves leathery, oblong, 14-24 cm long, nerves 11-15 pairs; 
cymes paniculate, 3-4 cm across; corolla fleshy. In forests; Bukit Timah (Md. 
Shah & Ali 4144), Bukit Mandal. 


Psych. helferiana Hook.f. 
Tiny shrub, densely hairy; leaves membranous, lanceolate or elliptic, 15-22 cm 
long; cymose heads globose; corolla white. In forests; Gardens’ Jungle, Chua Chu 
Kang, Bukit Timah (Md. Shah 749). 


Seed plants of Singapore 1X 161 


Psych. maingayi Hook.f. 
Slender climber, pubescent: leaves thick membranous, elliptic. narrowed at 
base, 5-9 cm long, nerves 5-6 pairs: cymes terminal and axillary; corolla yellowish 
green or white, 3-4 mm long. In woods near the sea: Kranji (Ridley 6924). Pasir 
Panjang. 


Psych. malayana Jack 
Shrub, to 2 m tall; leaves leathery, elliptic or elliptic-lanceolate, narrowed at 
base, 12-24 cm long, nerves 12-18 pairs; cymes corymbiform, 6-8 cm across. In 
edge of forests; Bukit Timah, Bukit Panjang (No specimens available). 


Psych. obovata Wall. 
Climbing shrub, glabrous; leaves ovate to obovate, leathery, 5-8 cm long. nerves 
5-6 pairs: corymbiform inflorescence large, many-flowered: corolla greenish 
white, 2-3 mm long, the tube very short, 5-lobed. In forest edges: Gardens’ 
Jungle, Kranyi (Ridley 2874). 


Psych. ovoidea Wall. 
Slender climber, hairy: leaves leathery, ovate or cordate, 4-6 cm long, nerves 5-7 
pairs: cymes dense, 2 cm across: corolla white. 4-lohed. In forests, Seletar, 
Gardens Jungle, Bajau (Ridley s.n. in 1894). 


Psych. penangensis Hook.f. 
Woody climber, glabrous: leaves fleshy, leathery, obovate-elliptic, 7-11 cm long. 
nerves 5-9 pairs; cymes 4-5 in corymbiform inflorescence. 8-10 cm across: corolla 
white. In forests; Gardens’ Jungle (Ridley s.n. in 1900); Water Catchment Areas. 


Psych. ridleyi K. & G. 
Low climber, 2 m tall, glabrous; leaves membranous, lanceolate to oblong- 
elliptic 10-14 cm long, nerves 8-10 pairs: branched cymes 4-5 cm across; corolla 
white. In forests; Gardens’ Jungle, Bukit limah (Ridley 10818). 


Psych. rostrata Bl. 
Erect shrub: leaves thin membranous, elliptic-oblong, narrowed at base, 7-14 cm 
long, nerves 5-6 pairs: cymes in small panicles: corolla white. In forests: Seletar. 
Pulau Ubin (No specimens available). 


Psych. sarmentosa BI. 
Climber. often creeping with aerial roots on tree trunks: leaves thin leathery, 
elliptic, 6-10 cm long, nerves 7-10 pairs; cymose corymbs 4-5 cm across: corolla 
greenish. In forests; Kranji, Serangoon, Changi (Ridley 2875). 


Psych. stipulacea Wall. ex Roxb. 
Stout low shrub, glabrous; leaves membranous, oblanceolate or elliptic, 12-22 cm 
long, nerves 10-12 pairs; cymes densely flowered, 1-1.5 cm across; corolla green- 
ish white. In forests; Bukit Timah (No specimens available). 


Randia anisophyllea (Jack ex Roxb.) Hook.f. (Alt. name: Porterandia anisophylla 
Ridl.) 
Small tree, to 10 m tall, often much branched, hairy: leaves obovate or elliptic, 
the base subequal, 10-30 cm long, nerves 10-14 pairs: cymes terminal, in dense 
clusters, 3-6 cm across; corolla tubular, 1.5-1.8 cm long, 5-lobed, white; berry 


162 Gard. Bull. Sing. 38(2) (1985) 


ovoid, 3-4 cm across, green, many-seeded, crowned by the 5-lobed calyx. In 
forests; common in Bukit Timah (Burkill HMB 318). (Randia as noted by Backer 
and Bakhuizen f. is an “extremely heterogenous” genus. The Malayan species 
were formerly dispersed in seven sections. Recently it was revised by Mr K. M. 
Wong (in Mal. Nat. J. Vol. 38). The names which appear in Mr Wong’s paper are 
cited here as alternative names). 


Randia auriculata (Wall.) Steud. (Alt. name: Aidia corymbosa (Bl.) Wong) 
Stout climber; leaves leathery, elliptic or oblong, 10-15 cm long, nerves 6-8 pairs; 
cymes in terminal and axillary corymbs, 6-8 cm across, hairy; berry globose, 5-6 
mm across, 2-loculate, many-seeded. In forests; Singapore (Wallich). 


Randia cochinchinensis (Lour.) Merr. (Alt. name, Aidia cochinchinensis Lour.) 
Shrub or small tree, to 12 m tall; leaves lanceolate to oblong-elliptic, 10-15 cm 
long; leathery; cymes in small dense clusters, axillary, 4-6 cm across, many- 
flowered; corolla 2-3 mm long. In forests; Chua Chu Kang (Ridley 5662). 


Randia fragrantissima Rid]. (Alt. name: Oxyceros fragrantissima (Ridl.) Wong) 
A stout climber, with paired recurved wood thorns, glabrous; leaves elliptic or 
ovate, leathery, 11-16 cm long; corymbiform inflorescences terminal and axil- 
lary, 4-5 cm across, of many fragrant flowers; corolla tubular, 2.5-3.5 cm long, 
creamy white. In forests, climbing on trees; Bukit Timah (Ridley 13022), Pulau 
Tekong, Changi. 


Randia longiflora Lam. (Alt.: name, Oxyceros longiflora (Lam.) Yamazak1) 
Climber with paired hooked thorns; leaves leathery, obovate or oblong, 4-8 cm 
long; umbellate cymes terminal or axillary; corolla 3-4 cm long, white. In tidal 
rivers and mangrove swamps; Changi, Pasir Ris (Furtado SFN 18640). 


Randia macrophylla R. Br. ex Hook.f. (Alt. name: Rothmannia macrophylla 
Bremek). 
Slender shrub, to about | m high, pubescent; leaves membranous, oblanceolate, 
18-30 cm long, subsessile, nerves 11-14 pairs; flowers 1 or 2, in upper axils, 
pendulous; corolla trumpet-shaped, 15-20 cm long, 8-10 cm across at the mouth, 
white with purple spots inside (“‘purple trumpet’’). In forests; Singapore (Hullett 
339). 


Randia macrantha DC. (Alt. name: Euclinia longiflora Salisb.) 
A garden shrub, 3-4 m tall, with a cluster of long (20-23 cm long) white flowers at 
the end of a branch. Native to tropical Africa, sometimes cultivated. 


Randia penangiana K. & G. (Alt. name: Oxyceros penangianus Tirvang.) 
Woody climber with stout paired axillary thorns (to 1 cm long); leaves elliptic, 
thin-leathery, 8-10 cm long, nerves 5-7 pairs; cymes 3-4-flowered, terminal and 
axillary; corolla 3-4 cm long, white. In forests; Gardens’ Jungle (Ridley s.n. in 
1902), Bukit Timah. 


Randia scandens (BI.) DC. (Alt. name: Oxyceros scandens (Bl.) Tirvang.) 
Climber, with paired recurved hooks, glabrous; leaves elliptic-oblong, thick 
leathery, 10-15 cm long; flowers 3 in a cyme; corolla tubular, 4-5 cm long, white. 
In forests, climbing on trees; Bukit Timah (Ridley s.n. in 1894). 


a 5 iti 


Seed plants of Singapore 1X 163 


Saprosma glomerulatum K. & G. 
Low shrub, glabrous, twigs and leaves foetid when bruised; leaves thin-leathery. 
elliptic or ovate, 10-16 cm long, nerves 7-9 pairs; flowers 4-5 in a sessile axillary 
cluster; corolla salver-shaped, 3-4 mm long, 4-lobed; fruit depressed globose, 
blue. In forests; Bukit Timah (Md. Shah & Samsuri 3896), Bukit Mandai. 


Scyphiphora hydrophyllacea Gaertn.f. 
Shrub, rarely a small tree, glabrous; twigs and petioles reddish when young; 
leaves ovate, leathery, upright, 3-5 cm long; flowers in dense cymes, 3-4 cm 
across; corolla about 1 cm long, cylindric, 4-lobed, pinkish; drupe green then 
whitish, 1 cm long, 6-8 grooved with two ribbed pyrenes inside. Common in 
mangroves and along muddy seashores; Jurong (Ridley s.n. in 1894). Vern. 
Cengan. 


Tarenna adpressa (King) Corner (= Stylocaryna adpressa King, Tarrena lancifoiia 
Rid.) 
Shrub, to 2 m tall; branches 4-angled, pubescent; leaves lanceolate or narrowly 
elliptic, thin-leathery, 15-20 cm long, nerves 6-10 pairs; many-flowered cymes 
forming dense corymbs, terminal; corolla cylindric, 1 cm long, 5-lobed; berry 
globose 5 mm across, whitish, with 2 pyrenes inside. In damp forests; Bukit 
Timah (Ridley s.n. in 1896). 


Taren. fragrans (BI.) K. & V. 
Shrub, to 2 m tall; leaves oblong or elliptic, 6-15 cm long; corymbs 5-7 cm 
across; berry globose, black. In forests; Geylang (Ridley 10933), Katong. 


Taren. grandifolia (Hook.f.) Ridl. 
Shrub, 1 m tall; leaves oblong-elliptic, thin-leathery, 10-20 cm long, nerves 8-12 
pairs; cymes 2-3 cm across, in corymbs, pubescent; corolla 1.2 cm long, white. In 
forests; Bukit Timah, Seletar (No specimens available). 


Taren. ridleyi (Pears.) Ridl. 
Shrub, 60 cm high, glabrous; leaves broad-elliptic, thin-leathery, 12-15 cm long, 
nerves 10-14 pairs; corymbs 5-6 cm across; berry fusiform. In open damp forests; 
Chua Chu Kang (Ridley s.n. in 1892). 


Taren. mollis (Wall.) Rid. 
Small tree, hairy; leaves lanceolate to ovate, leathery, 10-15 cm long, nerves 


10-12 pairs; corymbs 4-5 cm across; corolla white; berry black. In forests, Tanglin 
(Ridley s.n. in 1905). 


Taren. stellulata Rid]. 
Shrub, to 1 m tall; leaves elliptic, cuspidate, 10-20 cm long, nerves 7-10 pairs: 
corymbs 3-5 cm across; corolla 1 cm long; berry ovoid, 1 cm long. In forests; 
Woodlands (Ridley 11645); Bukit Mandai. 


Timonius compressicaulis (Miq.) Boerl. (= T. finlaysonianus Hook.f.) 
Shrub; leaves fleshy-leathery, oblanceolate or elliptic, 8-15 cm long, nerves 4-5 
pairs; flowers unisexual, on separate trees; male in short cymes about 2 cm long; 
corolla silky white, 2-3 mm long; female solitary; berry globose, bluntly 4-angled, 
many-seeded. In tidal mud along seashores; Serangoon (Ridley 2762). 


164 Gard. Bull. Sing. 38(2) (1985) 


Timon. flavescens (Jack) Baker (= T. peduncularis Ridl.) 
Small tree; leaves thin-leathery, elliptic, narrowed at both ends, 6-10 cm long, 
nerves 4-7 pairs; male cymes 3-7 flowered; corolla yellow, 1-1.2 cm long; berry 
oblong, 5-6 mm long, 4-angled, red. In open places; Bukit Mandai, Kranji, 
Seletar (Kadim Tassim 512). 


Timon. wallichianus (Korth.) Valeton 
Small tree; leaves lanceolate, 6-9 cm long, glabrous above, silvery hairy beneath, 
nerves 7-11 pairs; male cymes in dense clusters, 1.5-2.5 cm across; female cymes 
3-flowered, stalked; berry oblong, 1.2-1.5 cm long, bluntly 4-shouldered. In 
forest edges; Bukit Timah, Changi, Chua Chu Kang (Ridley s.n. in 1889). 


Timon. wrayi K. & G 
Tree, 15 m tall; leaves leathery, obovate-elliptic, base narrow, 12-20 cm long, 
nerves 6-7 pairs; male flowers 6-8 in a cyme, pubescent; berry solitary, ellipsoid 
or globose, crowned by calyx-lobes. Changi (No specimens available). 


Uncaria attenuata Korth. 
Slender climber; leaves elliptic, 6-12 cm long, nerves 6-8 pairs; flowers in a 
globose head, 3-4 cm across; peduncles 2-5 cm long, hairy; corolla slender, 8-10 
mm long; capsules fusiform, 2-valved, many-seeded. In forests; Bukit Mandai, 
Bukit Timah (Ridley s.n. in 1889). (Species of Uncaria are climbers, climbing by 
the aid of short, hook-shaped modified lateral branches. They can supply clear 
drinking water). 


Uncar. cordata (Lour.) Merr. (= U. pedicellata Roxb.) 
Climber; leaves coriaceous, ovate-elliptic, 8-12 cm long, nerves 7-8 pairs; heads 
4-5 cm across; peduncles 4-5 cm long; corolla slender, 1.5-2 cm long. In open 
forests; Bukit Timah, Tuas (Goodenough 2852). 


Uncar. gambir Roxb. 
Slender climber or bush (in cultivation); leaves ovate-oblong, 8-14 cm long, 
nerves 4-5 pairs; flower-heads 3-4.5 cm across; peduncles 2.5 cm long, slender, 
glabrous; corolla 1-1.2 cm long, tubular, slender, red, the lobes oblong, white. 
Native to Sumatra and Borneo, formerly cultivated in large plantations. An 
astringent extract, the gambier or pale catechu, prepared by boiling down the 
leaves and stems, is used by tanners and dyers. 


Uncar. glabrata (Bl.) DC. 
Slender climber, glabrous; leaves oblong, 6-8 cm long; flower-heads 2-2.5 cm 
across; peduncle thick, 2.5 cm long; corolla tubular, 6-8 mm long. In forests; 
Jurong, Bukit Mandai (Ridley 2846), Pulau Ubin. 


Uncar. jasminiflora Hook.f. 
Slender climber; leaves thin-leathery, elliptic, acuminate, 6-12 cm long, nerves 
5-6 pairs; flower-heads about 3 cm across; corolla slender, 1-1.2 cm long. In 
forests and mangrove swamps; Jurong, Bukit Mandai (Ridley 1041/5). 


Uncar. longiflora (Poir.) Merr. (= U. pteropoda Miq.) 
Large climber; leaves leathery, glabrous, elliptic, 12-16 cm long, nerves 7-8 pairs; 
petioles winged; flower-heads 2.5-3 cm across; peduncle as long. In forests; Bukit 
Timah (Ridley 2854), Gardens’ Jungle. 


i — = 


Seed plants of Singapore 1X 165 


Uncar. ovalifolia Roxb. 
Slender climber; leaves membranous, elliptic, shortly acuminate, 4-7 cm long, 
nerves 3-4 pairs; flower-heads 2-2.5 cm across; peduncle 1.5-2.5 cm long, slen- 
der; corolla tubular, 1.5 cm long. In damp forests; Chua Chu Kang, Ang Mo Kio 
(Ridley s.n. in 1889), Gardens’ Jungle. 


Uncar. roxburgiana Korth. 
Slender climber, soft hairy; leaves thin-leathery, ovate 5-8 cm long, nerves 5-6 
pairs; flower-heads 1.5-2 cm across; peduncles 1 cm long; corolla 1-1.2 cm long, 
slender, pinkish. In forest margins; Bukit Timah, Bukit Panjang, Chua Chu 
Kang (Ridley 6716). 


Uncar. sclerophylla Roxb. 
Large climber, covered with red soft hairs; leaves leathery, ovate or oblong, 
10-15 cm long, nerves 9-10 pairs; flower-heads 7-10 cm across; peduncle 6-8 cm 
long; corolla 2-3 cm long; silvery hairy. In thickets; Jurong, Gardens’ Jungle, 
Water Catchment Areas (Ridley 10635). 


Urophyllum glabrum Wall. ex Roxb. 
Slender shrub, to 2 m tall, glabrous; leaves thin-leathery, elliptic, acuminate, 
10-14 cm long, nerves 5-8 pairs; axillary clusters few-flowered, shortly pedun- 
cled; corolla greenish-yellow; berry globose, top flattened, orange, many- 
seeded. In forests; Ang Mo Kio (Ridley 61160). 


Uroph. griffithianum Hook.f. 
Shrub or small tree; leaves leathery, elliptic or oblong, 12-18 cm long, nerves 7-8 
pairs; axillary cymes many-flowered. In forests; Gardens’ Jungle, Changi, Mac- 
Ritchie Reservoir (Md. Shah & Md. Ali 3873). 


Uroph. hirsutum Hook.f. 
Shrub, or small tree, soft hairy; leaves oblong-lanceolate, 7-12 cm long, nerves 
8-10 pairs; flowers in small cymes; corolla 2-3 mm long, hairy. Common in 
forests; Gardens’ Jungle, Chua Chu Kang (Ridley 3906, ©’). 


Uroph. macrophyllum (BI.) Korth. 
Shrub or small tree; leaves elliptical oblong, acuminate, 3-8 cm long, nerves 9-11 
pairs; flowers in dense short-peduncled umbels, pubescent; corolla cylindric, 2-3 
mm long. In forests; Kranji. 


Uroph. streptopodium Wall. 
Slender shrub, to 2 m tall; young branches 4-angled, yellow, soft-hairy; leaves 
thin-leathery, elliptic-oblong, 5-15 cm long; flowers in short, dense cymes about 
1 cm long. In forests; Bukit Timah, Jurong, Pulau Ubin, Chua Chu Kang (Ridley 
4906). 


Uroph. trifurcum Pears. 
Shrub or small tree, glabrous; leaves leathery, oblong-elliptic, 12-18 cm long, 
nerves 10-12 pairs; flowers in small umbels, usually 3 umbels in a common 
peduncle. In forests; Changi, Kranji. 


Warszewiczia coccinea (Vahl.) Klotzsch. 
Shrub, 3-6 m tall; leaves oblong or obovate, 15-60 cm long; cymes in terminal 
panicles; one of the calyx-lobes of the outermost flowers enlarged, petaloid, 
elliptic, bright red, 3-11 cm long; corolla-tube yellow or orange, 1 cm long, 
5-lobed. Native to tropical America, sometimes cultivated in gardens. 


166 Gard. Bull. Sing. 38(2) (1985) 


126. CAPRIFOLIACEAE 


Key to the Genera 


A. Leaves pinnately or bipinnately compound; flowers regular, in terminal corymbose inflorescence .... 


sigavaun sd bie tnaere sens ace cn brussseeeeust@ae alt Ca@Myat ep Reeds sy othe yeas 1 ote ee ane Sambucus 
A. Leaves simple 

By) Flowers regular) in terminal panicles! .31.................cc.cssecaecesed een tee Viburnum 

B, Flowers irregular, in axillary pairs: visti 3) .di451) 08. SO ee eee Lonicera 


Lonicera japonica Thunb. 
Twining shrub; leaves simple, opposite, ovate, 3-7 cm long; flowers axillary, in 
pairs; corolla tubular, 2-lipped, 4-5 cm long, at first yellow, then becoming white, 
very fragrant; the bract below each flower-pair leaf-like, 1-1.8 cm long. Native to 
S. China and Japan. Another introduced species with smaller bracts is probably 
referable to L. confusa DC. of S. China. 


Sambucus javanica Reinw. ex BI. 
Erect shrub, 2-3 m tall; leaves simple pinnate; flowers small, white, fragrant, in 
a large terminal corymbose inflorescence. Native to Java, sometimes cultivated. 
Another species, S. canadensis L. from N. America, with bi-pinnate leaves, 
occasionally also cultivated. 


Viburnum sambucinum Bl. 
Large shrub; leaves lanceolate-elliptic, 12-20 cm long; flowers small, white; 
drupe red. Formerly found in Chua Chu Kang (Ridley 6829), Kranji, Bukit 
Panjang and Bukit Mandai, now probably extinct. 
127. CAMPANULACEAE 


Key to the Genera 


A. Flowers regular; anthers free; inflorescences axillary, in scorpioid cymes ................ Pentaphragma 
A. Flowers irregular; anthers mostly joined to form a tube 
B. Corolla-tube long and narrow, the limb not 2-lipped ...............ccececeeeeeeeneee eee enen es Laurentia 
B. Corolla-tube short, the limb .2-lipped tid) AR pe ARR ee eee Lobelia 


Laurentia longiflora (L.) Peterm. (= Isotoma longiflora (L.) Presl) 
Herb, 30-50 cm high, thick-stemmed, with milky sap; leaves simple, narrowly 
oblanceolate, 8-11 cm long, dentate or pinnatifid; flowers axillary, solitary; 
corolla white, tubulate, 7-11 cm long, 5-lobed near the top. Native to the West 
Indies, under hedges or along roadside. Vern. Star of Bethlehem. 


Lobelia zeylanica L. (= L. affinis Wall.) 
Ascending herb, to 15 cm long, usually pubescent; leaves ovate, acute, 1.5-4 cm 
long; flowers solitary, axillary; corolla white or pale blue, 2-lipped, upper lip 
2-lobed, lower 3-lobed, 6-8 mm long; capsule ribbed. In damp spots; Bukit 
Timah, Kranyji (Ridley s.n. in 1890). 


Pentaphragma horsfieldii (Mig.) Airy-Shaw (= P. scortechinii K. & G.) 
Succulent, woolly herb; leaves alternate, ovate-elliptic, slightly unequal-sided; 
flowers small, densely arranged in thick, curved scorpioid cymes, 5-6 cm long; 
corolla campanulate, yellowish; berry many-seeded. In damp forests, often on 
banks; formerly found in Bukit Timah, Sungei Morai, Pulau Damar (Ridley s.n. 
in 1894), Chua Chu Kang, now confined to Bukit Timah. 


Seed plants of Singapore LX 167 


Pent. ridleyi King 
Easily distinguished from the above species by its glabrous, often narrowly 
elliptic leaves with a narrowed base. In damp forests; Bukit Timah, Bajau 
(Burkill 723). According to Airy-Shaw, this is a natural hybrid, namely 
x elliptica Poulsen, a cross between P. acuminata Airy-Shaw and P. horsfieldii. 


128. GOODENIACEAE 


Scaevola taccada (Gaertn.) Roxb. (= S. frutescens Krause, S. sericea Vahl) 
Succulent shrub; leaves fleshy, oblong-obovate, round-tipped, 15-25 cm long, 
spirally arranged; flowers in leaf-axillary branched clusters; corolla white or 
tinged lilac, about 2 cm long, the tube split open along the upper side, limb 
5-lobed; drupe white, the endocarp corky, 1-2 seeded. A seashore plant, all 
round the coasts; Changi, Seletar, Labrador, Pulau Ubin, P. Tekong (Ridley s.n. 
in 1890). 


129. STYLIDIACEAE 


Stylidium tenellum Swartz (?) 
Slender shrub, 5-20 cm high, rarely branched; leaves obovate, alternate, 1-1.2 cm 
long; flowers very small, zygomorphic, purple, usually 2-3 together. In damp 
places in India and in the central and northern Malaya, recorded in Singapore by 
a specimen (Furtado s.n. in 1924) bought from a local Chinese drug shop. The 
identification of this specimen is highly questionable. 


130. COMPOSITAE 


Synoptic key to the genera* 


1. Flower-heads several in a cluster, surrounded by bracts 
Zo SO MEIS Ol OWel-HCaGs enanibe hel 0. Ais oh sgt fC Neri aged att Lees ec leeee Sphaeranthus 
2 Gas OF Newer tenUs 2 MAMeREO 22.1). /.eTai tee .te ress caese teres Elephantopus, Sparganophorus 
1. Flower-heads solitary or in branched inflorescences, not in clusters 
3. Leaves (at least the lower ones) opposite or crowded at the base of the stems 
4. Creeping or climbing herbs 
SO 1 ee lee eel aay eae eee Acanthospermum, Tridax, Wedelia 
i menerarem iin). 6 Ute treet i BO A. Eupatoria, Mikania 
4. Erect herbs 
6. Flower-heads in leaf-axils or at shoot-apex, sessile or nearly so, not in branched infloresc- 
REN Ss, ce ate, SE trek Be Vek Pe i, Sk Es hg Syke Eleutheranthera, Enydra, Synedrella 
Ana Peay oe etre Adenostemma, Ageratum, Bidens, 
Eclipta, Siegesbeckia, Spilanthes 
3. Leaves alternate; flower-heads usually in branched inflorescences 


OR ey epee) Re ees Poe ee ee ree eer Dicrocephala 
7. Flower-heads mostly cylindric, not globose 
8. Shrubs or shrubby, rarely trees ......................... Blumea (p.p.), Pluchea, Vernonia (p.p.) 
8. Herbs 
OE a ee Centipeda, Gynura, Microglossa 
Bo Breet Ob 38 Blumea (p.p.), Emilia, Erechtites, Erigeron, Vernonia (p.p.), Youngia 


Acanthospermum australe O. Ktze. (= A. brasilum Schrank) 
Greening herb, branched; leaves opposite, ovate, toothed, 1-2.5 cm long; flower- 
heads small, axillary; flowers white, the outer ones rayed. Native to tropical 
America, reported in Singapore in the last century. 


* Many cultivated genera are not included. 


168 Gard. Bull. Sing. 38(2) (1985) 


Adenostemma lavenia (L.) O. Ktze. (= A. viscosum Forst.) 
Erect shrub, to 1 m tall; leaves opposite, ovate, varying from 5 to 18 cm long, 
stalked and toothed; flower-heads in terminal branched and spreading infloresc- 
ences; flowers white, all tubular; achenes warty. In waste ground, often in damp 
places; Chua Chu Kang, Pulau Ubin (Hullett 78), Bukit Panjang. 


Ageratum conyzoildes L. 
Herb, erect and hairy, to 1 m tall; leaves opposite below and alternate above, 
ovate, 2-9 cm long, toothed; flower-heads usually 3-4 (each stalked) together on 
a common stalk in leaf-axils and terminal; flowers white or pale blue, all tubular; 
fruit black. Common in waste places throughout the Island, a garden weed. 


Artemisia lactiflora Wall. ex DC. 
Like the species below, also strongly aromatic, but leaves usually smaller (0.5-2.5 
cm long) and not white hairy beneath. Native of China, sometimes cultivated as a 
garden plant. 


Art. vulgaris L. 
Branched perennial, very aromatic; leaves ovate or lanceolate, 2-10 cm long, 
pinnately-lobed, densely white hairy beneath; flower-heads very small, in much 
branched panicles; light green, with one series of ray flowers enveloping the 
tubular ones. Native of north temperate countries, formerly commonly culti- 
vated for medicine in Chinese villages. 


Bidens pilosa L. 
A branched herb, 30-50 cm high; leaves opposite, simple or compound (with 3-5 
leaflets), 1-12 cm long, toothed; flower-heads on the top of branched stalks, each 
about 2 cm across, with marginal white or yellow ray-flowers; achenes long and 
narrow, black, bristled. Native of America, in waste places, common in hill 
stations in Malaya, occasionally recorded in Singapore. 


Blumea balsamifera (L.) DC. 
Shrubby herb, up to 3 meters or more high, hairy, aromatic; leaves alternate. 
elliptic, toothed, 8-40 cm long, the stalk often with 2-3 pairs of narrow-lobed 
appendages; flower-heads numerous, forming a very large terminal and axillary 
branched inflorescence; flowers all tubular, yellow. In open places, Bajau 
(Goodenough 2742). Leaves and stems with a strong smell of camphor when 
crushed; used in local medicine. Vern. Sumbong, Ngai camphor. 


Blumea lacera (Burm. f.) DC. 
Erect herb, 30-50 cm tall, strongly smelling; leaves oblong, 2-5 cm long, the 
lower leaves often lobed; flowers yellow. A weed on roadsides and grassland. 


Centipeda minima (L.) A. Br. & Asch. 
Tiny prostrate herb, much branched; leaves oblong, alternate, pinnately lobed, 
0.4-2 cm long; flower-heads solitary, axillary, very small (0.2-0.4 cm across); 
marginal flowers white or purplish, central disc-flowers yellow. In waste ground; 
Botanic Gardens, Woodlands (Sinclair SF 39237). 


Chrysanthemum morifolium Ramat. (= C. indicum Hort.) 
The chrysanthemum is imported as a pot plant or a cut flower, very various in 
form and colour in the flower-heads. Native of E. Asia of ancient cultivation. 


3B BITE 


Seed plants of Singapore LX 169 


Coreopsis lanceolata L. 
Perennial, 30-60 cm tall, branched; leaves tufted at base, opposite above, narrow 
spoon-shaped entire or 2-3 lobed; flower-heads 5-6 cm across, long stalked, ray 
and central flowers yellow. Native to N. America, sometimes cultivated. 


Coreopsis tinctoria Nut. 
Like the above, but leaves twice-pinnately divided; flower-heads 3-6 cm across; 
ray flowers reddish brown and central flowers reddish purple. Native to N. 
America. 


Cosmos sulphureus Cav. 
Tall herb, to 1 m high; leaves opposite, finely divided; flower-heads 4-6 cm 
across, solitary or few together, on long stalks; ray flowers light to deep yellow or 
orange, central ones yellow. Native to Mexico, sometimes cultivated. 


Crassocephalum crepidioides (Benth.) S. Moore 
Erect herb, fragrant, hairy, to 1 m high; leaves oblong or elliptic, 8-15 cm long, 
pinnately lobed or pinnatifid; flower-heads cylindric, 1.5 cm long, yellow with a 
reddish top, forming small branched corymbs. A weed of tropical African origin, 
a fairly recent introduction. 


Dahlia pinnata Cav. 
Perennial herb with tuberous roots; leaves opposite, simple or pinnately divided; 
flower-heads flat to globose; marginal ray flowers white, pink to puple, flat, 
tubular or rolled; central tubular flowers yellow. Native to Mexico, sometimes 
cultivated. 


Eclipta prostrata L. (= E. alba Hassk.) 
Branched creeping herb; leaves opposite, lanceolate, 2-10 cm long, almost 
sessile; flower-heads stalked (the stalk 2-5 cm long), solitary or 2-3 in a leaf axil, 
less than 1 cm across; ray flowers white. Common weed in waste ground. 


Elephantopus scaber L. 
Herb, varying from 2-3 cm to over 30 cm high, hairy; leaves often crowded at the 
base of stem, lanceolate or oblong, 4-15 (or more) cm long; flower-heads 
few-flowered, several heads together in a bracteate cluster at the end of branched 
terminal inflorescence; flowers all tubular, pinkish white. A common weed often 
found in waste places throughout the Island. Vern. Tutup Bumi. 


Eleutheranthera ruderalis (Sw.) Sch.-Bip. 
Annual aromatic herb, 10-60 cm high; leaves ovate, 1.5-7 cm long; flower-heads 
terminal and axillary, single or in pairs, few-flowered; flowers yellow. In sandy 
places; Changi, Pulau Ubin (Furtado 18629). 


Emilia sonchifolia (L.) DC. 
Erect shrub, 20-30 cm tall; leaves alternate, from rounded (lower ones) to 
heart-shaped (upper ones), toothed or variously lobed; the upper leaves sessile, 
the lower ones stalked; flower heads narrow-tubular, 1.5-2 cm long, solitary or 
few in long-stalked terminal inflorescence; flowers all tubular, upper part bright 
pink. A common weed in waste ground and in gardens. Vern. Katumbi Jantan. 


170 Gard. Bull. Sing. 38(2) (1985) 


Enydra fluctuans Lour. 
Creeping herb; leaves opposite, narrowly oblong, 2-10 cm long, entire or tooth- 
ed, fleshy; flower-heads white or greenish, axillary, less than 1.5 cm across, 
sessile. In wet places; Geylang (Ridley 10829). Leaves and stem are edible (called 
‘Buffalo spinach’). (The generic name is often incorrectly spelled as Enhydra). 


Erechtites hieracifolia (L.) Rafin. ex DC. 
Erect shrub, to 1 m tall; leaves alternate, narrowly oblong, 10-12 cm long, 
toothed; flower-heads cylindric, 1-1.5 cm long, 2-5 together in branched terminal 
(or upper axillary) inflorescence; flowers tubular, yellow, with silky-white pap- 
pus hairs. In waste grounds; native to tropical America. 


Erecht. valerianifolia (Wolf) DC. 
Differs from the above species in the broader and often deeply lobed leaves and 
in the pinkish-orange flowers with reddish violet pappus hairs. In waste grounds; 
also native to tropical America. 


Erigeron sumatrense Retz. 
Annual, to 2 m high, branched; leaves alternate, long-spoon-shaped, 3-14 cm 
long, the upper ones smaller and narrower than the lower ones; flower-heads 
bell-shaped, about 0.5 cm long, greenish-white, numerous in large and much 
branched inflorescence. A common weed in waste and cultivated ground. Vern. 
Sumbong Jantan. 


Eupatorium odoratum L. f. 
Tall herb sometimes shrubby; leaves opposite, ovate, pointed, 5-10 cm long; 
flower-heads pale blue to white, about 1.5 cm long, in paniculate inflorescences; 
flowers all tubular. A weed originally from N. America, arrived from Thailand to 
Malaya during the first World War, hence the Malay name “‘Pokok German”’; 
occasionally found in Singapore since late 1970s. 


Gaillardia pulchella Fong. 
Annual, 15-30 cm tall; leaves alternate, narrowly spatula-shaped, 5-12 cm long, 
entire or wavy-lobed; flower-heads 5-10 cm across, long-stalked; ray flowers 
10-18, wedge-shaped, yellow or red; central flowers purplish. Native to N. 
America, sometimes cultivated. 


Gerbera jamesonii Bolus. ex Hook. 
Perennial, hairy; leaves pinnately lobed, 20-30 cm long, in a basal rosette; 
flower-heads solitary, arising from a long stalk; ray flowers narrow, red or 
orange, in one or two rows surrounding the central white flowers. Native of 
Transvaal, S. Africa, often planted. 


Gynura procumbens (Lour.) Merr. (= G. sarmentosa DC.) 
Creeping or climbing herb; leaves alternate, fleshy, ovate, pointed, 5-8 cm long, 
toothed; flower-heads tubulate, 1.5-2 cm long, 3-7 together in terminal branched 
inflorescence; flowers all tubular, orange yellow in a purple involucre. Common 
in open places or in secondary growth; Bukit Mandai, Bukit Timah (Ridley s.n. 
in 1891), Tanglin. Vern. Akar Subiak. 


Helianthus angustifolius L. 
Herb, branched, 1 m or so high; leaves linear or linear-lanceolate, 4-30 cm long; 
flower-heads 5-7 cm across, several to many (in short and long stalks) in a leafy 
panicle; ray flowers golden yellow, surrounding a dark purple centre. Native to 
N. America, sometimes growing in borders or for cut flowers. 


Seed plants of Singapore 1X 171 


Helianthus annuus L. 
Stout annual, 1-2 m tall; leaves alternate, ovate, 7-45 cm long, serrate; flower- 
heads 15-35 cm across, often nodding and turning towards the sun (thus ‘Sun 
Flower’); ray flowers yellow, surrounding a brown-purple centre; achenes 
(known as ‘Sunflower seeds’) large, edible. Native of N. America, sometimes 
cultivated. |i  # 


Helianthus tuberosus L. 
Erect shrub, 2-4 m high; tuberous root ellipsoid, up to 15 cm long (‘Jerusalem 
artichoke’); leaves ovate oblong, 10-20 cm long; flower-heads 5-8 cm across, in 
leaf panicles; ray flowers golden yellow, surrounding a bright yellow centre. 
Native of N. America, sometimes cultivated for its edible tubers. 


Lactuca indica L. 
Herb, 1-2 cm high; leaves lanceolate, 8-25 cm long, entire or dentate; flower- 
heads in large panicles; branches of the panicle with a few linear bracts only at 
the base. Native to E. Asia, cultivated as vegetables. Ht) , ‘| # 


Lact. sativa L. 
Herb, 0.3-1 m high; leaves broadly oblong, 6-14 cm long, finely dissected; 
flower-heads in large flat-topped inflorescence with many cordate bracts. Native 
to Europe, sometimes cultivated. 


Melapodium divaricatum (Pers.) DC. 
Herb with angular stems; leaves opposite, ovate, 2-9 cm long; flower-heads 
solitary, axillary, 1-1.5 cm across; ray flowers yellow. Native to Central America, 
sometimes cultivated as a pot plant. 


Mikania cordata (Burm. f.) B. L. Robins. 
Fast-growing climbing herb; leaves cordate or ovate, tip acuminate, base heart- 
shaped, 3-12 cm long; petiole 1-8 cm long; flower-heads cylindric, 6-9 mm long, 
in dense corymbs; corolla white or pale yellowish, all tubular; bracts enveloping 
the individual florets 5-6 mm long. Formerly very common in waste lands; 
Tanglin (Burkill 407); now almost completely replaced by the following slightly 
smaller-flowered exotic species, M. micrantha. This species sometimes was 
erroneously called M. scandens (L.) Willd., a species restricted to N. America. 


Mikania micrantha HBK. 
A very fast-growing climbing herb, commonly called ‘Mile-a-minute’; leaves 
narrowly or broadly ovate, tip pointed, base heart-shaped, 4-8 cm long, 2-4 cm 
wide; flower-heads small, white, usually hanging at the ends of a much branched 
inflorescence; bracts enveloping the individual floret 3-4 mm long. In forest 
edges, secondary growth and open places, common.* 


Pluchea indica (L.) Less. 
Shrubby herb, to 1.5 m tall; leaves spirally alternate, thick, oblong, 2-6 cm long, 
toothed short-stalked; flower-heads cylindric, in much branched terminal in- 
florescences; flowers all tubular, purplish or white. Common in tidal swamps and 
on sea coast, rarely in inland grounds; Changi, Geylang, Jurong (Ridley s.n. in 
1888). Vern. Pokok Beluntas. 


* Information on this species was kindly supplied by Dr Richard Corlett. 


172 Gard. Bull. Sing. 38(2) (1985) 


Rudbeckia serotina Nutt. 
Hairy herb, to | m high; leaves alternate, lanceolate or oblong, 5-12 cm long; 
flower-heads solitary, terminal, 10-12 cm across; ray flowers golden yellow, 
surrounding a dull brown or black centre (thus called ‘black-eyed Susan’). Native 
to N. America, sometimes growing in gardens. 


Sigesbeckia orientalis L. 
Hispid herb, 0.3-1 m tall, branched; lower leaves petiolate, ovate or oblong 5-20 
cm long; upper leaves narrower, smaller and sessile; flower-heads greenish, 1.5-2 
cm across, in loose panicles; the ray flowers surrounded by linear glandulate 
spreading bracts. In moist waste places, not common. 


Solidago altissima L. 
Herb, to 1 m tall; lower leaves forming a rosette at the base; the upper ones 
alternate, spatulate, 3-14 cm long; flower-heads golden yellow, rather small, 
usually borne on one side of the branches (hence called ‘Golden rod’) of a large 
panicle. Native to N. America; this and some other related species sometimes 
sold as cut flowers. 


Sparganophorus vaillantii Crantz. 
Fleshy herb, 30-40 cm high; leaves alternate, elliptic, 2-12 cm long; flower-heads 
sessile in leaf-axils, flattend, 0.5-1.5 cm across; flowers white, all tubular. In 
damp places; a tropical American weed. 


Sphaeranthus africanus L. 
Branched herb; stems and branches winged; leaves alternate, oblong, 2-3 cm 
long; flower-heads globose, less than 1 cm across, consisting of many small 
tubular florets, greenish white, solitary, terminal or in upper axillary, shortly 
stalked. In muddy waste places or in ditches; Geylang (Ridley 5069). 


Spilanthes acmella (L.) Murr. 
Herb to 50 cm tall; leaves opposite, broadly ovate, 3-6 cm long, toothed; 
flower-heads terminal, solitary, ovoid, about 1 cm across, long-stalked (12 to 15 
cm long); marginal ray-flowers yellow; achenes black. In waste ground; Seletar 
(Hullett 630); formerly sold in local herb shops for curing toothache (thus “the 
toothache plant’’). Vern. Krabo. 


Syndrella nodiflora (L.) Gaertn. 
Annual, with branched slender stems; leaves opposite, elliptic, 1-5 cm long; 
flower-heads axillary, yellow, small and few-flowered, sessile or subsessile, fruit 
black. Native of tropical America. (Similar to Eleutheranthera ruderalis but 
differing from the latter in having narrower stalked flower-heads with smaller 
bracts and 2-3 stiff spines on the achenes). 


Tagetes erecta L. 
Herb, 30-50 cm tall; leaves opposite (lower ones) or alternate (upper ones), 5-12 
cm long, deeply lobed or compound; flower-heads solitary, terminal, yellow or 
orange, 5-10 cm across; ray flowers many, flat or rolled. Native to Mexico, but 
horticulturally known as African Marigold. Another species also from Mexico 
but erroneously called French marigold, Tagetes patula L. having smaller flower- 
heads (3-4 cm across), the ray flowers are marked with red colour. 


Seed plants of Singapore 1X | Ae 


Tithonia diversifolia Gray 
Shrubby; leaves alternate, ovate, entire or 3-5 lobed, 10-25 cm long; flower- 
heads 8-10 cm across; ray flowers (about 12) orange yellow, surrounding the 
small yellow central tubular flowers. Native to Mexico and C. America, some- 
times cultivated. 


Tridax procumbens L. 
Creeping herb; leaves opposite, narrowly ovate, coarsely serrate, 1-5 cm long; 
flower-heads 2 cm across, on upright long-stalk (10-30 cm long); ray flowers pale 
yellow to white. A weed in open dry, sandy places; native to Central America. 


Vernonia arborea Buch. Ham. (incl. V. javanica DC.) 

Tree, 10-20 m tall, much branched above; leaves alternate, ovate to oblong, 8-20 
cm long; flower-heads 0.5 cm long, white or pink, 5-6 flowered, all tubular; 
paniculate inflorescences terminal, widely branched. In mature secondary 
forests; Changi, Chua Chu Kang, Jurong and Water Catchment Areas (Corner 
s.n. in 1936). Vern. Merambong. (This is the only tree species of the Compositae 
in this region. Two forms can be recognized: one a tall straight tree with glabrous 
leaves, and the other often stunted and gnarled, with pubescent leaves. The 
latter is either treated as a variety, 1.e., var. javanica Clarke, or a separate 
species, V. javanica DC.). 


Vernonia cinerea (L.) Less. 
Herb, to 1 m tall; leaves alternate, generally ovate, 1-8 cm long, but very 
variable; flower-heads narrowly cylindric, 6-7 mm long, violet or pink, all tubu- 
lar, many in a terminal, much branched inflorescence. Common weed, in waste 
places or in gardens. 


Vernonia patula (Dryand.) Merr. (= V. chinensis Less.) 
Similar to V. cinerea, but stouter and with larger flower-heads and bracts. 
Technically, the two aspects can be distinguished by the following characters: in 
V. patula, achenes 5-angular, pappus hairs in | series; in V. cinerea, achenes 
cylindric, pappus hairs in 2 series. Common in waste grounds and clearings; 
Geylang (Teruya 2523). 


Wedelia biflora (L.) DC. 
Hairy scrambling herb, sometimes shrubby; leaves ovate, pointed, 3-nerved, 
3-15 cm long; flower-heads usually solitary, bright yellow, about 1.5 cm across, 
with ray-flowers surrounding the tubular ones. On sandy beaches; Pulau Ubin 
(Hullett 387). Vern. Serenai Laut. 


Wed. trilobata (L.) Hitch. 
Creeping herb; leaves fleshy, oval-shaped, toothed or 3-lobed; flower-heads of 
prominent yellow ray-flowers. A species of tropical American origin, often 
planted in gardens as a ground cover. 


Xanthium inequilaterum DC. (=X. strumarium Auct. non L.) 
Herb, 30-60 cm tall; leaves alternate, ovate, toothed and lobed, 5-10 cm long; 
flower-heads unisexual, in spikes on upper leaf-axils; achenes oblong, enclosed 
in the enlarged involucre covered with hooked bristles known as ‘burs’. Formerly 
recorded once at Rochore (Ridley s.n. in 1904) as a casual weed, now apparently 
disappeared. 


174 Gard. Bull. Sing. 38(2) (1985) 


Youngia japonica (L.) DC. (= Crepis japonica (L.) Benth.) 
Herb, 10-30 cm high; leaves nearly all basal, 5-10 cm long, round-topped, 
toothed or lobed below; flower-heads oblong, 5 mm long, yellow tipped, in 
loosely branched inflorescences. A weed commonly found in hill stations in 
Malaya, occasionally reported from Singapore. 


Zinnia linearis Benth. 
Herb, 20-60 cm tall; leaves linear or lanceolate, 1-6 cm long; flower-heads 2-3.5 
cm across; ray-flowers orange. Native of Mexico, sometimes cultivated. 


Zinnia elegans Jacq. 
Herb, 0.3-1 m tall, branched; leaves elliptic, ovate, 3-15 cm long; flower-heads 
terminal, 3-7 cm across; ray-flowers red, pink, white or yellow. Native to Mex- 
icO, sometimes cultivated. 


Micropropagation of Lagerstroemia speciosa (L.) Pers. (Lythraceae) 


LIM-HO CHEE LEN and LEE SING KonG 


Botanic Gardens, Singapore 


Abstract 


Lagerstroemia speciosa (L) Pers., a common road-side tree of Singapore, was successfully mass- 
produced using the tissue culture technique. Nodal segments were the best explants as they could 
produce more multiple shoots than do shoot tips with BAP and 2ip treatments. The excised shoots were 
rooted in agar medium or sterilised sand, supplemented with IBA. The former was found superior. 
About 90% of the plantlets survived when transplanted to soil. 


Introduction 


Lagerstroemia speciosa is commonly planted along roadsides and in open spaces 
throughout Singapore for its shade. These trees also produce colourful flowers in 
abundance. There is some variation in the flowering characteristics of some of the 
trees in the population established. Some of them consistently flower more profuse- 
ly than other trees located in close proximity. As it is the intention of the Parks 
and Recreation Department to introduce colours to our landscape effectively, it 
would be beneficial to plant avenues and groves of those trees recognised for 
their superior flowering qualities. A row of trees with uniform flowering habits is 
most desirable. To achieve this objective, there is a need to clone trees having 
the above characteristics. Although this could be achieved through rooted stem 
cuttings, the number that can be propagated is limited. Thus, the use of the tissue 
culture technique to mass propagate these trees is investigated and this paper 
reports the results of in vitro propagation of Lagerstroemia speciosa. 


Materials and Methods 


Young shoots from the crowns of mature trees as well as from the basal sprouts 
of tree stumps were collected. The leaves were removed and the axils were brushed 
lightly to remove dirt. This was carefully done so as not to break or damage the 
axillary buds at the nodes. The axillary buds were prominent and had an average 
length of 2-3 mm. Shoot tips and nodal segments with buds were separated, washed 
in tap water and used as explants. The length of the explants, i.e., shoot tips and 
nodal segments used were 5 mm and 10 mm respectively. They were then surtace- 
sterilised by dipping in 98% ethanol for 5-10 seconds, and then in 30% clorox 
solution with Tween 80, for 45 minutes to 1 hour. The sterilised materials were 
rinsed five times in sterile water. 


The basal medium consisted of Murashige and Skoog (1962) salt solution at the 
following concentrations: — a) macronutrient at half and, b) micronutrient at full 
strength, c) 3% sucrose. To test the response of explants, the media were further 
supplemented with IAA (Indole 3-acetic acid) and Kinetin (6 furfurylaminopur- 
ine). The concentrations used for both IAA and Kinetin ranged from 0-10.0 ppm, 
used either individually or in combination with one another. These concentrations 
were tested in 7 X 7 factorial combinations. There were 49 treatments altogether 
and each treatment was replicated 10 times. 


176 Gard. Bull. Sing. 38(2) (1985) 


For multiple shoot induction, media containing BAP (6-benzylaminopurine at 0, 
1, 2, 3, 4 mg/l) or a combination of BAP at 1 mg/l and 2ip (N6 — [A° isopentyl] - 
adenine at 1, 2, 3 mg/l) were used. Ten replicates of each treatment were prepared. 
After eight weeks of growth, shoot proliferation was evaluated in terms of increase 
in fresh weight, number of shoots formed and average length of shoots. The pH of 
the media was adjusted to 5.5 before autoclaving. The culture tubes were auto- 
claved at 120°C at 1.5 kg/cm ~* pressure for 20 minutes. The cultures were kept 
under 1 klx lighting provided by true-lite tube on a 12 hour photoperiod. The 
environment temperature was kept at 21-25°C. 


Results and observations 
The various explants responded differently when cultured on the various media. 
The pattern of response to the various treatments by each of the explants tested, 


are described below. 


Growth of Nodal Segments and Shoot Tips 


Table 1. Growth response of Lagerstroemia speciosa on MS salt with 49 combina- 
tions of [AA and Kinetin. 


Callus, shoot, root development were observed in 20-day old cultures. 
C : slight callus 


CC: vigorous growth of callus 
CCC : abundant callus 

S : shoot 

— : no growth 


RR: root 


Micropropagation of Lagerstroemia speciosa 177 


Of the 2 types of explants, nodal segments responded better and Kinetin had 
little effect on the growth (Table 1). Satisfactory callus and shoot formation were 
obtained with IAA at 0.5-2 mg/l. There was no significant difference in the 
response among the various combinations of Kinetin & IAA used. An average of 2 
shoots developed from the axils of the nodal segment (Plate 1). Growth was first 
observed 7 days after inoculation, and about 20 days thereafter, shoots of 3-5 cm 
length were obtained. In most of the media, an initial swelling at the basal portion 
of the explant was observed. The swelling was due to cell divisions in the cortical 
layers leading to an increase of the tissue volume. After about 2 weeks, masses of 
friable and white callus were formed at the base of the culture shoots. Shoot tips on 
the other hand, usually browned off or showed less growth and they normally grew 
into single shoots (Plate 2). 


Since nodal segments gave best growth, they were used as explants for all 
subsequent experiments. 


Shoot multiplication 


Multiple shoots developed within 15 days after inoculation. An average of about 
3 buds were seen at each node and these developed into shoots in about 2 to 5 
weeks. The best results were obtained in BAP medium at 1 mg/l (Table 2) and 
shoots reached a length of 1.8 cm. Increasing the BAP concentration promoted the 
growth of callus but caused a reduction in the shoot length (Table 2). The leaf 
blades were very much reduced, thus resulting in cane-like structures. There was no 
significant difference in the number of shoots formed. 


Table 2. Effect of BAP on the induction of multiple shoots. 


Concentration of BAP Average shoot 
(mg/l) Average no. of shoots length 


* The shoots were counted in 30-day old cultures. 


With the addition of 2ip into the medium, the multiplication of shoots improved 
markedly (Table 3). The average number of shoots increased three-fold to 9 
shoots per explant. The average shoot length also increased from 1.8 to 3.2 cm and 
there was no significant difference at different concentrations of 2ip (1-3 mg/l). All 
the cultures showed healthy growth with a high frequency of normal leaves and 
shoots with long internodes (Plate 3). It was possible to mass-produce the shoots at 
this stage. Shoots which had elongated to about 5-10 nodes could be excised and cut 


178 Gard. Bull. Sing. 38(2) (1985) 


into several nodal segments for further multiplication on the same medium. In 
these segments, the axillary buds enlarged and additional young shoots appeared in 
about 2 weeks (Plate 4). In about 6 weeks, they multiplied again (Plate 5) and 
could be excised for rooting. This process could be repeated to produce the 
required number of plantlets. 


Table 3. Effect of combination of 1 mg/l BAP and 2ip (1-3 mg/l) on the induction of 
multiple shoots. 


* 


Average fresh 
weight of shoots 


Average shoot 


Concentration of 2ip Average no. of 
length 


(mg/l) shoots 


* The shoots were counted in 30-day old cultures. 


Rooting of excised shoots 


Tests were conducted on excised shoots of 3 to 4 cm length planted onto basal 
medium supplemented with IBA (Indole-3-butyric-acid at 0, 2, 3, 6 mg/l) or in 
sterile sand supplemented with MS salt solution and IBA (at 5, 10, 20, 40 and 60 
mg/l). In each of the treatments, there were 10 replicates. 


No root was formed in the control medium but with IBA added into the medium, 
healthy roots were formed (Plate 6). The average number of roots and the average 
root length did not vary with different IBA concentrations (Table 4). The most 
important effect of IBA was to induce roots and rooting percentage increased 
proportionally with the concentration until it reached 100% at 6 mg/l. At low IBA 
concentration, soft callus formed at the base of the tissues. 


Table 4. Effect of different concentrations of IBA on rooting of shoots on MS agar 
medium. : 


* 


Concentration of BAP Average length Rooting 


Average no. of root Percentage 
of roots 


* The roots were counted in 30-day old cultures. 


Micropropagation of Lagerstroemia speciosa 179 


= 


Table 5. Effect of different concentrations of IBA on rooting of shoots in sand 
medium, supplemented with MS salt. 


* 


Concentration of 2ip 


Average length Rooting 
of root Percentage 


Average no. 
of roots 


“ The roots were counted in 30-day old cultures. 


For rooting in sand supplemented with different concentrations of IBA, the best 
results were obtained at the concentrations of 10-20 mg/l (Table 5). Although more 
roots were formed, they were short (Plate 7). Only about 50% of the cultures 
produced roots at the optimal IBA concentrations of 10-20 mg/l. 


After rooting, the small plantlets were removed from the media and potted with 
either a mixture of coco-peat and sand (1:1) or a mixture of sand, coco-peat and soil 
(1:1:3). The potted plantlets were covered with a plastic sheet for about 2 weeks to 
maintain high humidity. Water stress is thus reduced and the development of 
epicuticular wax allowed (Grant & Aston, 1977). After 2 weeks to 1 month, the 
plants had hardened and the plastic sheets were removed. Ninety percent of the 
plantlets which rooted from agar media survived whereas only 50% of the plantlets 
rooted directly from sand survived. They all resembled plants produced by cuttings 
(Plate 8) and were similar in growth habit and form. However, whether rooted in 
agar media or sand, the potting-out survival rate was observed to decrease sharply 
for plantlets kept in the rooting culture for longer than two months. 


Discussion 


Tissue culture techniques provide viable alternative methods of mass-production 
of healthy plants with uniform characteristics. These techniques have been success- 
fully applied to many ornamental plants such as orchids (Lim-Ho, 1982), herbs and 
shrubs (George & Sherrington, 1984). Although it is known that the explants of 
woody and tree species of Gymnosperms and Angiosperms are difficult to grow in 
vitro and to induce regeneration, methods for micropropagation of temperate and 
tropical trees have been developed from a variety of explant sources during the last 
few years (Bonga & Durzan, 1982; Dodds, 1983). 


180 Gard. Bull. Sing. 38(2) (1985) 


In this study we have demonstrated the feasibility of mass-production of L. 
speciosa using nodal segment and shoot tip cultures from stump sprouts. The ability 
of explants from stump sprouts to regenerate shoots in vitro is an advantage as 
plantlets produced would resemble the mother plants in terms of growth character- 
istics and habits. This would be especially useful when the main objective is to 
clone trees identified for their profuse flowering characteristics. The results of the 
present study also support the view that tissues at the basal region of a tree trunk 
still remain juvenile irrespective of the age of the tree (Bonga & Durzan, 1982; 
Vietiez et. al., 1983; 1985). 


The most commonly used cytokinin for shoot multiplication in trees is BAP and 
its effectiveness used singly has been well demonstrated in the seedling tissues of 
Calophyllum, Eugenia, Fagraea (Lee & Rao 1980, 1982), Swietenia (Rao & Lee, 
1982), and mature tissues of Quercus robur (Vietiz, 1985) and Eucalyptus (Gupta 
et. al., 1981) from in vitro. BAP alone was not effective in promoting shoot 
multiplication of L. speciosa. The addition of 2ip induced a higher multiplication 
rate, showing an interaction effect of BAP and 2ip on shoot multiplication. Such 
results are very rare in literature (Bonga & Durzan, 1982). Further studies could be 
initiated on the interaction effects of other cytokinins with BAP to gauge the 
response of shoot multiplication. 


Rooting experiments in agar media were more successful than those tested on 
sand. Furthermore, when the plantlets were potted out, there was 90% survival 
rate. Further work would help to ascertain the best rooting media that would 
encourage better survival of plantlets when potted out. The present methods also 
show that it is possible to rapidly produce large numbers of plantlets of Lager- 
stroemia in vitro and it is suggested that this method is available for the propagation 
of other useful trees. 


Acknowledgements 


The authors wish to thank Mrs Quek-Phua Lek Kheng and Mrs Teo-Lee Guek 
Choon for their competent assistance in the experiments. 


Literature Cited 


Bonga, J.M. and D.J. Durzan (1982). Tissue Culture in Forestry. Martimus Nijhoff/ 
Dr. W. Junk Publications. The Hague. 


Dodds, J.B. (1983). Tissue Culture of Trees. The Avi Publishing Comp. Inc. 
Weatport, Connecticut, USA. 


George, E.F. and P.D. Sherrington (1984). Plant Propagation by Tissue Culture. 
Exegetics Ltd., Eversley, Basingstoke, England. 


Grout, J.M. and D.C. Anston (1977). Transplanting of cauliflower plants regener- 
ated from meristem culture, water loss and water transfer related to changes in 
leaf wax and xylem regeneration Hort. Res., 17: 1-7. 


Gupta, P.K., A.F. Mascarenhas and V. Jajannathan (1981). Clonal propagation 


of mature trees of Eucalyptus citriodora Hook, by tissue culture. Plant Sci. Letter 
20: 195-201. 


Micropropagation of Lagerstroemia speciosa 181 


Lee, S.K. and A.N. Rao (1980). Tissue culture of certain tropical trees. In Plant 
Cell Cultures: Results and perspectives (Sala. F. and other Eds.). Elsevier 
North Holland Biomedical Press. Amsterdam, 305-311. 


& ____. (1982). In-vitro plantlet development in tropical trees — 
Calophyllum inophyllum and Eugenia grandis. Tissue Culture of Economically 
Important Plants (A.N. Rao ed.) COSTED Publ. Singapore 185-190. 


Lim-Ho, C.L. (1982). Tissue culture of local orchid hybrids at the Singapore 
Botanic Gardens. Tissue Culture of Economically Important Plants (A.N. Rao 
ed.) COSTED Publ. Singapore 295-300. 


Murashige, T. and F. Skoog (1902). A revised medium for rapid growth and 
bioassays with tobacco culture. Plant Physiol. 15: 473-497. 


Rao. A.N. and S.K. Lee (1982). Importance of tissue culture in tree propagation — 
Proc. 5th Intl. Cong. Plant Tissue & Cell Culture. (Akio Fuji Wara ed.) 
Tokyo, Japan. 


Vieitez, A.M., A. Ballester, M.L. Vieitez and E. Vieitez (1983). Jn vitro plantlet 
regeneration of mature chestnut — Jour. Hort. Sci. 58: 457-63. 


Vieitez, A.M., M. Carmen San-Jose and E. Vieitez (1985). Jn vitro plantlet 
regeneration from juvenile and mature Quercus robur, L. Jour. Hort. Sci. 60: 
99-106. 


Plate 1. Two shoots developed from lateral bud Plate 2. Single shoot developed from shoot-tip 
(20-day old culture). (20-day old culture). 


Metbienhn shits 


Cd 2 i ae 


Plate 3. Multiple shoots on MS supplement with BAP (1 mg/l) + 2ip (1-3 mg/l) (18-day old culture). 


182 


: 
; 


Plate 4. Young shoots arising Plate 5. Multiple shoots from nodal segments (40-day old culture). 
from nodal segment 


(14-day old culture). 


SSS 
SS 
SS 
S 
SS 

SS 


roots (30-day old culture). 


Plate 6. Lagerstroemia rooted in agar medium showed less but longer 


Plate 7. Lagerstroemia rooted in sand showed more but shorter roots (30-day old culture). 


Plate 8. Well-established Lagerstroemia plantlets produced by tissue culture (60 days in planung 
media). 


184 


A New Account of the Genus Horsfieldia (Myristicaceae), Pt 3* 


W.J.J.O. de WILDE 
Rijksherbarium, Leiden, The Netherlands 


EFFECTIVE PUBLICATION DATE: 15 FEB. 1986 


Contents 


Enumeration and description of the species 46-70 .........-..seeee eee ee eee eee eect ene ee nent ees page 185-225 


46. Horsfieldia sabulosa Sinclair Fig. 1B(46); 22 


Horsfieldia sabulosa Sinclair, Gard. Bull. Sing. 27 (1974) 133 — Type: Sinclair & Kadim 10491 (K; iso: 
BA. BL ENY, SAR, 4.0% 


Tree 10-37 m. Twigs stout, terete, neither ridged nor lined but with wart-like 
thickenings marking the leaf-scars, (3-)5-10(-20) mm diam., grey-brown, non- 
striate, rather late glabrescent from grey-brown to rusty tomentum with hairs 
Q-5-1.0(-1.5) mm long; bark lower down grey-blackish, usually longitudinally crack- 
ing and often + flaking; lenticels inconspicuous. Leaves in 3-5 rows, generally 
bunched towards the apex of the twigs, coriaceous, elliptic-oblong to (sub) lan- 
ceolate, nearly parallel-sided or broadest + at the middle, 9-21 x 2-6 cm, base 
attenuate, tip acute (to short-acuminate); upper surface glabrous, + shining or not, 
olivaceous to brown, lower surface rather pale brown, early glabrescent, with 
scattered dark brown to blackish dots, these roundish or sometimes elliptic or 
line-shaped; midrib flat above, glabrous; nerves 12-20 pairs, flat above and below, 
the lateral arches and the tertiary venation invisible; petioles long in proportion to 
the blades, 25-50 x 2-4 mm, rather late glabrescent, tomentum with hairs 0.5-1.5 
mm; leaf bud rather short and stout, 10-15 x (3-)4-5 mm, with dense grey-brown to 
rusty tomentum with hairs 0.5-1.5 mm long. Inflorescences behind the leaves on 
older twigs of 15-20 mm thick, emerging from the axils of the rough and woody 
wart-like petiole-scars on the older bark, densely pubescent with hairs 1.0-1.5 mm 
long; in & (submature seen on San 15146): 4-5 x 2-3 cm, c. 2-3 times ramified, 
many-flowered, flowers in loose clusters of 5-10 each; common peduncle c. 15 mm 
long; inflorescences + few-flowered, 3-5 cm long; bracts elliptic to lanceolate, + 
boat-shaped, 4-15 mm long, densely pubescent with hairs 0.5-1.5 mm but glabrous 
inside, late caducous; flowers 3-(or 4-) valved, perianth glabrous, pedicels glab- 
rous, at base articulate. Male perianth (immature) globose or depressed- globose, 
c. 0.7 X 1.0 mm; pedicel c. 0.8-1.0 mm long; perianth (in bud) with valve-sutures to 
c. Y2-way, buds not collapsing on drying; valves c. 0.2 mm thick. Androecium 
(immature) + globose c. 0.4 x 0.5-0.6 mm, = circular in transverse section; anthers 
c. 12-15, + completely sessile, apical cavity small; androphore narrow, + 0.2 mm 
long. Female flowers (according to Sinclair, l.c., p. 134): ovary immature, glabrous. 
Fruits 1-6 per infructescence, ovoid, top and base broadly rounded, 3.0-5.0 x 
2.5-4.0 cm, glabrous, drying dark brown to somewhat bluish black, not warted: 
pericarp hard, 5-8(-10) mm thick; stalk 2-8 mm long; perianth not persisting. 


Distribution. Borneo: Sarawak (Serian Dist., Bintulu Dist., Mulu Nat. Park), 
Brunei, Sabah (Sipitang). 


*Continued from Gdns’ Bull. Sing. 38 (1): 144. 


186 Gard. Bull. Sing. 38(2) (1985) 


BORNEO. Sarawak: Brunig 956; (Chai) S. 39647; Sinclair & Kadim 10248 — Brunei: BRUN 579, 
0828; Sinclair & Kadim 10437, 10491 — Sabah: San. 15146, 17560. 


Ecology. A moderate to large tree of mixed forest on sandy or peaty soil, 
deep yellow sands, sandy loam, or heavy yellow clay soil; in Agathis forest, ridge 
forest; 0-100 m alt. Flowers May & June, fruits May to December. 


Vernacular name. Kumpang-perawan (Iban). 
NOTES 


1. Fieldnotes. Tall tree, buttresses absent. Bark dark grey, bark of trunk longitu- 
dinally fissured. Sap red, copious. Inner bark reddish brown, laminated, fibrous; 
sapwood soft, pinkish, with a hollow centre c. 2.5 cm wide. Leaves glossy above, 
very glaucous beneath, midrib greenish yeilow. Fruits yellow, inside apricot; rami- 
florous with many fruits on each branch. 


2. A-very characteristic species because of its thick branches with bunched 
leaves, the leaves with long petioles in (3-) 5 rows, ramiflorous. Sinclair discussed 
its relationship with H. wallichii, which has resembling fruits. This has also almost 
similar dark brown dots on the lower leaf surface. 


3. According to Koster and Baas (1981, p. 152) the leaves have the unique 
feature of having an 1so-bilateral mesophyll. 


47. Horsfieldia atjehensis de Wilde, sp. nov. Fig. 1B(47) 
Horsfieldia amygdalina auct. non (Wall.) Warb.: Merrill, Contr. Arn. Arb. 8 (1934) 61. 


Folia ramulorum fertilium sparsa, subtus punctis sparsis fusco-brunneis non-traumaticis induta (ut in H. 
glabra), cortice ramulorum delapso. — Type: W. & C.M. Bangham 882 (K; iso: A, NY, n.v.). 


Tree c. 10 m. Twigs terete, not ridged, towards the top 3.5-5(-8) mm diam., 
rather pale grey-brown to yellowish brown, early glabrescent, tomentum grey- 
brown with hairs less than 0.1 mm, lower down with the bark coarsely striate and 
tending to flake; lenticels rather conspicuous towards the top of the twig. Leaves in 
3-5 rows, thinly chartaceous, obovate-oblong or elliptic-oblong to oblong- 
lanceolate, broadest slightly above the middle, 13-25 x 4.5-9 cm, base long- 
attenuate, tip acute-acuminate; upper surface drying dark brown; lower surface 
early glabrescent, provided with regularly scattered brown to blackish non- 
traumatic larger dots (lens!); midrib flat above; nerves 10-12 pairs, flat above, 
marginal arches indistinct; tertiary venation forming a lax network indistinct or 
invisible on both surfaces; petioles 12-15 x 2.5-3.5 mm; leaf bud densely grey- 
brown pubescent with hairs less than 0.1 mm, moderately slender, c. 15 x 3.5-4 
mm. Inflorescences situated behind the leaves, very thinly pubescent to glabres- 
cent, hairs less than 0.1 mm; in ©: c. 3 times ramified, rather many-flowered, 7-14 
x 4-10 cm, common peduncle 10-20 mm long; @ inflorescences not seen; bracts 
elliptic-oblong, finely pubescent, 2-4 mm, caducous. Flowers 3-valved, in the male 
in loose clusters of 4-8 each, glabrous; pedicels glabrous, at base inarticulated. 
Male perianth (slightly immature) globose, c. 1.5 mm diam., at anthesis cleft to c. 
’2-way,; valves c. 0.2(-0.3) mm thick; pedicel slender, 1-1.5 mm long. Androecium 
subglobose to short-ellipsoid, c. 1.2 x 1.0 mm, top broadly rounded, in transverse 
section subcircular; anthers 11, almost completely sessile, free apices c. 0.1(-0.2) 
mm, curved over and + into the rather narrow apical cavity c. 0.3 mm deep; 


Fig. 22. Horsfieldia sabulosa Sinclair. 
a. shoot apex with densely bunched leaves with dispersed phyllotaxis, tomentum of young 
leaves partially fallen, x 12; b. older wood with leaf scars in dispersed phyllotaxis and 
immature male inflorescences, note bracts, X %2; c. smaller bract, <x 12; d. immature male 
flower, lateral view, X 12; e. ditto, longitudinally opened, showing androecium, x 12; f. 
longitudinal section of androecium, schematic, x 12; g. old wood with infructescence, fruits 
mature, seeds completely covered by aril, x 2. —a-f. from San. 15146; g. from BRUN 0828. 


columm broad, spongy-solid; androphore narrow, c. 0.2(-0.3) mm long. Female 
flowers and fruits not seen. 


Distribution. Known from only one collection in N. Aceh, Sumatra. 


Ecology. Mountainous forest at c. 1300 m (3500-5000 ft.); male flowers, still 
immature in January. 


187 


188 Gard. Bull. Sing. 38(2) (1985) 


NOTES 
1. Fieldnotes. Leaves leathery, glabrous, flower buds green. 


2. This is in many respects much related to and + intermediate between H. 
amygdalina, H. glauca, H. macrothyrsa, and H. sparsa, but still markedly distinct 
from all these species. 


H. amygdalina, a species from continental SE. Asia and the Andaman Isls. (not 
known from the Nicobar Isl. and Malaya), differs in general habit and further by 
the absence of blackish dots on the lower leaf surface and by the darker colour of 
the dried twigs. 


H. sparsa from lowland Peninsular Thailand, Malaya, and Sumatra, has a rather 
similar general habit, with a similarly pale bark on the twigs, but differs by the 
absence of blackish dots on the lower leaf surface, and by the flowers which have a 
rather different androecium with less anthers (7-9), and a generally longer 
androphore. 


H. macrothyrsa, occurring in N. and C. Sumatra, also in mountainous forest, has 
punctate leaves, but differs in the distichous phyllotaxis, and the much larger male 
flowers with a different androecium. 


Hitherto H. glabra (var. glabra) is found on Sumatra only in the lowland in the 
southern part, and all specimens from that area strongly differ in general habit and 
have the leaves distichous. The present new species has in common with H. glabra 
the punctate leaves but differs in various characters viz. a stouter habit, the leaves 
in 3-5 rows along the twigs, the bark rather pale and coarsely striate, tending to 
flake, and probably in the slightly smaller male perianths. Furthermore, it was 
found rather distant from the area of H. glabra at a much higher altitude. 


3. The Kew-specimen was identified by Sinclair as H. sucosa in 1959; in his 
treatment of 1975 it is enumerated under H. bracteosa. In the publication on the 
Bangham collections by Merrill (1934) the collection was treated under H. amygda- 
lina. 


48. Horsfieldia sucosa (King) Warb. Fig. 1B(48) 


Myristica sucosa King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 301, pl. 172 — Horsfieldia sucosa (King) 
Warb., Mon. Myrist. (1897) 322; Sinclair, Gard. Bull. Sing. 16 (1958) 416 (p.p., incl. lectotype, excl. 
fig. 45 = H. sparsa), pl. XII A; 28 (1975), 139 p.p. — Syntype: King’s Coll. 4078, 4647 (fr., lecto: K, 
L; CAL, G, n.v.; see notes), 10475; Wray 467; Maingay (Kew Dist.) 1300. 

H. bracteosa Henderson, Gard. Bull. Str. Settl. 7, 2 (1933) 120, pl. 30; Sinclair, Gard. Bull. Sing. 16 
(1958) 419, fig. 46 — H. bracteosa var. bracteosa: Sinclair, Gard. Bull. Sing. 28 (1975) 18. — Type: 
Henderson SFN 24521 (SING, n.v.; iso: K; DD, n.v.). 


Tree 6-20 m. Twigs terete, neither lined nor ridged, towards the apex 2-10 mm 
diam., pale grey-brown or straw-coloured, contrasting with the blackish colour of 
the dried petioles, (very) early glabrescent, tomentum rusty to greyish, hairs 
0.1-0.3 mm long; lower down the bark rather coarsely striate or not, with a 
tendency to flake, lenticels rather conspicuous only on the very young parts. Leaves 
in 2 or 3 rows, membranous to thin-chartaceous, elliptic-oblong to oblong-(ob) 
lanceolate, broadest at or above the middle, 14-28 x 4.5-8.5 cm, base attenuate or 
long-attenuate, tip acute-acuminate; upper surface glabrous, drying greenish- 
brown to blackish-brown, often with a greenish tinge, lower surface early glabres- 


New account of Horsfieldia 3 189 


cent, drying light brown, without larger blackish dots; midrib flat above or only 
slightly raised, glabrous; nerves 13-17 pairs, slender above, flat or slightly raised, 
lateral arches indistinct; tertiary venation forming a lax network indistinct or 
invisible above; petioles 10-20 x 2-3 mm, early glabrescent; leaf bud c. 10-15 x 2-3 
mm, with rusty to greyish brown tomentum of hairs 0.1-0.3 mm long. Inflor- 
escences situated behind the leaves, thinly pubescent or late glabrescent from hairs 
0.1-0.2 mm long; in CG: 3 or 4 times ramified, many-flowered, rather lax or 
condensed, 7-19 x 5-16 cm, common peduncle 10-20 mm long, flowers in clusters 
of 3-7 each; Q inflorescences rather few-flowered, 1-2 cm long; bracts + ovate-ellip- 
tic to lanceolate, acutish, pubescent, 1.5-4 mm long, rather late caducous; flowers 
either mostly 2-valved (Borneo, see further under the subspecies) or mostly 3- 
valved (Malaya, Sumatra), perianth glabrous, pedicels usually glabrous (see under 
subsp. sucosa), at base distinctly articulate and contrasting with the pubescent 
branches of the inflorescence. Male perianth globose to depressed globose, top 
flattish to broadly rounded, base rounded or broadly rounded, 1.2-1.5 x 1.5-2.0 
mm; pedicel 1.0-2.0 mm long, slender; perianth at anthesis cleft from c. 3 to 
(nearly) ¥2-way, on drying not or only slightly collapsed at the top, valves 0.2-0.3 
mm thick. Androecium depressed globose, 0.3-0.6 X 0.8-1.2 mm, top flattish or 
broadly rounded, circular or (in Borneo) broadly ellipsoid in transverse section; 
anthers 7-11, almost completely sessile towards the incurved apex, column broad, 
+ saucer-shaped, with a broad and flattish, rather shallow apical cavity reaching up 
to nearly ¥2-way deep; androphore rather narrow, 0.1-0.3 mm long. Female 
perianth (only seen in var. bifissa): broadly ellipsoid, 2.8-3.5 x 2.2-3.0 mm, 
2-valved, cleft to c. %4, valves 0.3-0.4 mm thick, pedicels 1-1.5 mm long, when 
young with minute hairs 0.1 mm long or less towards the base, ovary c. 1.5 mm 
diam., glabrous, stigma shallowly 2-lobed, c. 0.1 mm high. Fruits 1-4 per infructesc- 
ence, broadly ovoid-ellipsoid, top + narrowly rounded, 2.3-3.5 x 2.0-2.5 cm, 
glabrous, drying blackish with finely granulate surface, sometimes + tuberculate; 
pericarp c. 4 mm thick; stalk 1-2 mm long; perianth in Malayan specimens persis- 
tent (see further under the subspecies). 


Distribution. Malaya, Sumatra, Borneo. 


NOTE. Divided into two geographically separated taxa, mainly based on the 
differing number of valves of the perianth, a character regarded as important in the 
genus and on which, grosso modo, the genus can roughly be divided into two 
divisions. It is noteworthy that this character, here on a subspecific level, occurs 
within the division of species with mainly 3-valved perianths. 


KEY TO THE SUBSPECIES 


la. Perianths predominantly 3 (or 4)-valved. Malaya, Sumatra ..................c0ceeeeeee ees a. subsp. sucosa 
b. Perianths predominantly 2-valved. Borneo .................ccccceecueceeceecuecneneeeeeneeees b. subsp. bifissa 
a. subsp. sucosa Fig. 1B(48) 


Perianths predominantly 3-valved, rarely a few 2- or 4-valved; mature male 
perianths in bud 1.2-1.5 x 1.5-2.0 mm; androecium 0.4-0.6 x 0.8-1.2 mm; 
anthers 7-9 (Malaya) or 9-11 (Sumatra). Fruits 2.5-3.5 x 2.0-2.5 cm with persis- 
tent 3-lobed perianth. 


Distribution. Malaya, Sumatra. 


MALAYA (Perak, Kelantan, Trengganu, Pahang, Selangor, Malacca, Johore): FR/J 1252, 4212, 
5228, 14088, 14331, 15219, 15603; Kep. FN. 97901; King’s Coll. 4647, 10475; Phyt. survey Kuala 
Lumpur (Millard) 1825; Maingay 2422; SFN (Sinclair & Kiah) 39937, 40621, 40629. 


190 Gard. Bull. Sing. 38(2) (1985) 


SUMATRA (Jambi, Palembang, Benkulu): b.b. 1799, 31936; Kostermans 12020; Lambach 1311; 
Marsden (Hb. Hooker) s.n.; Roos & Franken 1704. 


Ecology. Dry land and seasonal swamp forest; recorded from sandy soils or 
sandstone; also disturbed forest; 0-500 m alt. Flowers and fruits throughout the 
year. 


Vernacular names. Merampat (Tamuan, Malaya); Pérédah bésar (Palembang 
Sumatra). ¢ 


NOTES. 


1. Fieldnotes. Bark smooth or shallowly fissured or thin-scaly. Slash bark lami- 
nated, reddish, with sticky reddish exudate. Wood yellowish or pink. Flowers 
yellow-green or yellow, scentless. Fruits shiny green, turning yellow or yellow-pink 
or pink-red. Aril orange-red. 


2. Dry fruits, over 3.5 cm long, have never been measured, but fresh fruits are 
recorded as 5-7.5 cm long, with a thick pericarp. 


3. As pointed out by Sinclair (1958, p. 418; 1975, p. 140) King’s syntype is 
heterogeneous. Only King’s Coll. 4647 and Maingay 1300 (also numbered 2422) 
belong to the present species. King’s Coll. 4647 (in fruit) is herewith designated as 
lectotype. 


4. King describes the flowers as externally pubescent; however, as explained by 
Sinclair (1958, p. 418) that material was very heterogeneous, and most likely this 
record is erroneous. The flowers of H. sucosa, as circumscribed presently, are 
generally glabrous, but the female flowers of Kostermans 12012 (Palembang) have 
pedicels somewhat pubescent towards the base. 


5. In the herbarium, most specimens of our present H. sucosa subsp. sucosa 
were initially identified by Sinclair as H. sucosa, a species accepted besides the 
related H. bracteosa. Later on, most of these specimens were re-identified by 
Sinclair as H. bracteosa; he indicated he was aware of the heterogeneity of his H. 
sucosa, from which I have presently segregated the new species H. sparsa. Sinclair’s 
lectotype-specimens of King’s material, King’s Coll. 4647 and 10475, however, are 
clearly identical with the type of H. bracteosa. 


b. subsp. bifissa de Wilde, subsp. nov. 


Perianthia mascula depresso-globosa. 1.2-1.4 x 1.5-1.7 mm, praecipue 2-valvata; androecium depresso- 
globosum, 0.3-0.4 x 0.8-0.9 mm, antheris 7. Fructus 2-2.5 cm longi, eius perianthium non-persistens. 
— Type: Kutei, Soegeng 58 (L; iso: BO, n.v.). 


Perianths predominantly 2-valved, the odd 3-valved perianth present; mature 
male perianths in bud 1.2-1.4 x 1.5-1.7 mm; androecium 0.3-0.4 x 0.8-0.9 mm; 
anthers 7. Fruits c. 20-25 x 20 mm, the perianth + caducous, 2-lobed. 


Distribution. Borneo: E. Sabah, E. and S. Kalimantan. 


BORNEO, E. Sabah: Elmer 21607; San. 68452, 73922 (NT 169). — E. & S. Kalimantan: Afriastini 
195; Endert 3213; Kostermans 8081, 9973; Soegeng Reksodihardjo 59; Hubert Winkler 2373, 2695. 


New account of Horsfieldia 3 19] 


Ecology. Lowland forest: on sandy loam soil, sandy ridge in Shorea laevifolia 
forest; 0-800 m. Flowers and fruits throughout the year. 


NOTES 


1. Fieldnotes. Bark smooth, undulately fissured; inner bark c. 10 mm thick. 
laminated, reddish-brown or yellow-brown; wood whitish or pale brown, or red- 
dish. Flowers yellow, the males very fragrant. 


2. Specimens ot the present new subspecies were formerly determined by Sin- 
clair as H. sucosa var. microcarpa, a name later on (1975, p. 20) published as H. 
bracteosa var. microcarya Sinclair. Other specimens belonging to Sinclair’s var. 
microcarya, incl. the type, are presently referred to a new species H. pallidicaula. 


3. H. sucosa is vegetatively very similar to H. pallidicaula and H. sterilis, the two 
being markedly different in the male flowers: these have a differently built 
androecium, and the pedicels are inarticulate at the base. 


4. It seems as if the distributional area excludes that of the related and resemb- 
ling species H. pallidicaula. 


49. Horsfieldia pallidicaula de Wilde, sp. nov. Fig. 1B(49) 


Cortex ramulorum pallidus, cinereus. non fusco-brunneus. Folia disticho vel sparsa. subtus non- 
punctata. Perianthia mascula subglobosa, 1.5-2 mm diam., 3-vel 4-valvatis: pedicelli graciles, 1-2 mm 
longi, basi non-articulati. Fructus late ellipsoidei, in sicco nigrescentes, perianthio persistente. — 
Type: Jacobs 5413 (L: iso: K; SAR, G. US, CANB, B.S, SING, n.v.). 


Tree 7-20 m. Twigs terete, neither lined nor ridged, towards the apex 2-10 mm 
diam., pale, whitish or greyish brown, contrasting with the blackish colour of the 
dry petioles (or petioles often greyish in the lower half), early glabrescent, tomen- 
tum with rusty or greybrown hairs 0.1-0.3 mm long; the bark lower down finely 
striate or not, slightly flaking or not, lenticels generally inconspicuous or absent. 
Leaves either in 2, 3 or 5 rows, membranous, oblong to oblong-lanceolate, 
broadest at or above the middle, 10-30 x 4-9.5 cm, base long-attenuate, tip 
acute-acuminate; upper surface drying brown to blackish-brown, lower surface 
drying brown to grey-brown, without larger blackish dots, early glabrescent: midrib 
flat or slightly raised above, glabrous; nerves 10-18 pairs, slender above, flat or 
slightly raised, lateral arches not distinct; tertiary venation forming a lax network. 
indistinct above; petioles 10-25 x 1.5-3 mm, early glabrescent; leaf bud rather stout 
to slender, 7-10 x 2-4 mm, covered by brown-grey rusty tomentum of hairs (0.1-) 
0.2(-0.3) mm long. Inflorescences situated generally behind the leaves, glabrescent, 
tomentum weak, with hairs 0.1-0.2 mm; in CG: (2 or) 3 times ramified, many- 
flowered, 3-9 x 2.5-7 cm, common peduncle up to 7 mm long (hence inflorescences 
often also ramified from near the base), the flowers more or less in clusters of 2-4: 
inflorescences rather few-flowered, 1-3 cm long; bracts lanceolate, acute, finely 
pubescent, 1-3 mm long, caducous; perianths 3- or 4-valved, in Q sometimes 
2-valved, glabrous, pedicel glabrous, at base inarticulate. Male perianth globose or 
broadly obovoid, 1.5-2.0 (-2.2) mm diam., top (broadly) rounded, base rounded to 
short-attenuate; pedicel 1-2 mm, slender; perianth at anthesis cleft to c. /3 to nearly 
¥2-way deep, not or but slightly collapsing on drying, valves 0.2-0.3 mm thick. 
Androecium short-ellipsoid to somewhat depressed-globose, 0.8-1.2 * 0.8-1.2 mm, 
top rounded, circular in cross-section; anthers 8-10, completely sessile (i.e., free 
apices 0-0.1 mm long only), incurved towards the apex of the androecium; column 


192 Gard. Bull. Sing. 38(2) (1985) 


broad and solid with a small and narrow apical cavity, 0.1-0.2 mm deep only; 
androphore narrow, short, 0-0.2 mm long. Female perianth broadly ellipsoid or 
subglobose, c. 2.5-3.0 X 2.5 mm, cleft at anthesis to ¥%4-¥3, valves (0.2-) 0.3 mm 
thick, pedicels 1-1.5 mm long, ovary subglobose to broadly ovoid, 1.7-2.0 x 1.5-2.0 
mm, glabrous, stigma shallowly 2-lobed, lobes 0.1-0.2 mm long. Fruits 1-5 per 
infructescence, broadly ellipsoid, top and base rounded, 1.5-4.0 x 1.0-3.0 cm, 
glabrous, drying blackish, finely granulate and sometimes tubercled, pericarp 
various; stalk 1-3 mm long; perianth persisting under the fruit (always?) (see 
further under the varieties). 


Distribution. Three varieties in Borneo (Sabah, Sarawak). 


KEY TO THE VARIETIES 


la. Fruits 15-17 x 10-12 mm; pericarp 1-1.5 mm thick. Leaves up to 23 cm long ...... b. var. microcarya 
b.) Frurts larger rk. 0 onion la sd ee SR Sa ro EA ae 2 
2a. Fruits c. 20-22 x 18 mm; pericarp 2-3 mm thick. Leaves up to 25 cm long .......... a. var. pallidicaula 


b. Fruits 37-40 x 27-30 mm; pericarp 6-8(-10) mm thick. Leaves 25-30 cm long ...... c. var. macrocarya 
a. var. pallidicaula Fig. 1B(49) 


Leaves 10-25 cm long, up to 7.5 cm wide. Fruits c. 20-22 x 18 mm; pericarp 2-3 
mm thick; stalk and perianth not seen. 


Distribution. Borneo: Sarawak, West Sabah, West Kalimantan (doubtful, see 
notes). 


BORNEO. Sarawak (1st, 3rd Div., Baram Dist.): Jacobs 5413; Hose 29, 86; S. 14971, 37692, — 
Sabah (West): SAN. 15254, 76783, 80505. — West Kalimantan: Hans Winkler 1435 (doubtful). 


Ecology. Primary lowland and lower mountainous forest, recorded from sand- 
stone; 0-700 m alt. Flowers throughout the year, fruits in June. 


Vernacular name. Kumpang balau (Iban). 
NOTES 


1. Fieldnotes. Bark smooth, dark grey; sapwood white-red. Perianth yellow or 
yellow-green, androecium pale pink-yellow, pollen white. 


2. The three varieties presently recognized are sympatric and are very similar in 
the vegetative characters. They are mainly distinguished by the remarkable differ- 
ences in fruit-size, and hence I am not quite sure to which variety the male 
flowering material belongs. 


The only known fruiting specimen regarded as belonging to the type-variety is 
San. 15254, with fruits of c. 21 * 18 mm and of which the calyx is lost. 


Also, of the only two known female flowering specimens one cannot be sure to 
what variety they belong for the same reasons. 


3. All three varieties of the presently proposed new species are vegetatively also 
very similar to the related species H. sucosa and H. sterilis, but both these differ 
essentially in the male flowers. 


New account of Horsfieldia 3 193 


4. Most specimens of our present vars. pallidicaula and microcarya were at first 
identified by Sinclair on the sheets as H. sucosa var. microcarpa, and in 1975 
published as Horsfieldia bracteosa var. microcarya. 


5. A noteworthy deviating specimen. The collection Hans Winkler 1435, from 
West Kalimantan, Sungei Bika, 5.1.1925, with mature male flowers, keys out 
together with H. pallidicaula. It deviates in general habit by its very thin membra- 
nous leaves which dry greenish. The inflorescences are rather weak, the flowers 
small, slightly longer than broad c. 1.5 x 1.4 mm, pedicel c. 1-1.5 mm, perianth- 
valves 3, thin, splitting the bud to nearly 2-way, the androecium is subglobose, c. 
0.8-0.9 x 0.7-0.8 mm, = sessile, anthers c. 9, sessile, the central-apical cavity being 
narrow and c. 0.2 mm deep. The specimen was collected in a marshy forest at c. 50 
m alt. Possibly it represents a separate taxon. The specimen was determined by 
Sinclair as H. carnosa, which it certainly is not. 


b. var. microcarya (Sinclair) de Wilde, comb. nov. 


Horsfieldia bracteosa Henderson var. microcarya Sinclair, Gard. Bull. Sing. 28 (1975) 20 — Type: Wood 
& Kapis San. 1697] (K; iso: L; KEP, SAN SING, n.v.). 


Leaves to 20 cm long, up to 7.5 cm wide. Fruits c. 15-17 x 10-12 mm; pericarp 
1-1.5 mm thick; stalk c. 2 mm long, persistent perianth 3-lobed. 


Distribution. Borneo: West Sabah, possibly E. Kalimantan. 


BORNEO. West Sabah: Wood & Kapis San. 16791. 
Ecology. Lowland forest at c. 100 m. Fruits in July. 


NOTE. Known only from the type, a collection which vegetatively completely fits 
the species generally, but distinct in its particularly small fruits with persistent 
3-lobed perianths. Possibly here belongs Leigton 943 (E. Kutei; fruits not seen). 


c. var. macrocarya de Wilde, var. nov. 


Differt a var. pallidicaula fructibus siccis c. 4 cm longis, pericarpio sicco 6-8(-10) mm crasso. — Type: 
Ding Hou 474 (L). 


Leaves 23-30 x 6-9.5 cm. Fruits 35-40 x 25-30 mm; pericarp 6-8(-10) mm thick; 
stalk 1-2 mm long, perisistent perianth 2- or 3-lobed. 


Distribution. Borneo: Sarawak (4th and Sth. Division) 


BORNEO. Sarawak: Ding Hou 474; S$ 32299. 


Ecology. Lowland mixed Dipterocarp forest; yellow sandy clay soil; 30-200 m alt. 
Fruits in July and October. 


NOTES 


1. Fieldnotes. Bark recorded as smooth to slightly flaky, or as longitudinally 
fissured. Fruit pink, aril pink. 


2. Known only from the type, with mature fruits, and one collection (S$ 32299) 
with immature fruit. Both have persistent perianths. These in the type are 
apparently 2-lobed, the ones of S$ 32299 are 3-lobed. 


194 Gard. Bull. Sing. 38(2) (1985) 


50. Horsfieldia sparsa de Wilde, sp. nov. Fig. 1B(50) 


Horsfieldia sucosa auct. non (King) Warb.: Sinclair, Gard. Bull. Sing. 16 (1958) 416, fig. 45, plate XII 
A; 28 (1975) 139. 


Cortex ramulorum pallidus, cinereus, non fusco-brunneus. Folia sparsa, subtus sine punctis brunneis 
non-traumaticis. Perianthium masculum subglobosum, c. 2 mm diam, 3-5 valvatum, pedicello basi 
non-articulato. Androecium depresso-globosum, 0.8-1.2 mm diam., antheris 7-9. Fructus ellipsoidei 
3-5.5 cm longi, in sicco nigrescenti, perianthio non-persistenti. — Type: Malaya, Ogata Kep. 110406 
(1, ISG; 35), 


Tree 15-40 m. Twigs terete, neither lined nor ridged, towards apex 4-11(-15) mm 
diam., pale whitish brown or grey-brown, contrasting with the brown-black colour 
of the dried petioles, early glabrescent, tomentum brown or grey-brown, hairs 
0.1-0.2 mm; bark lower down rather smooth or coarsely striate, sometimes with a 
tendency to flake, lenticels present but conspicuous only towards the apex. Leaves 
crowded or not towards the apex, arranged in 3-5 rows, thickly membranous to 
chartaceous, oblong to oblong-lanceolate, broadest at or above the middle, (10-) 
18-24 x 3.5-7 cm, base long-attenuate, top acute-acuminate; upper surface drying 
bright brown to dark brown, lower surface drying bright brown, glabrous (early 
glabrescent), without larger blackish dots; midrib above flat or slightly raised, 
glabrous; nerves 12-16 pairs, above slender, flat or slightly raised, lateral arches not 
distinct; tertiary venation forming a lax network, not or hardly visible above; 
petioles (12-)14-34 x (1.5-)2-3 mm; leaf bud rather stout, relatively short and 
broad, c. 8-12 x 3-5 mm, with dense tomentum of hairs c. 0.1-0.2 mm long. 
Inflorescence behind the leaves, subglabrous or early glabrescent, tomentum very 
weak, of hairs c. 0.1-0.2 mm; in CG: 2 or 3 times ramified, many-flowered, 6-12 X 4-6 
cm, common peduncle (2-)5-15 mm long, the flowers + fasiculate in clusters of 4-10 
each; 9-inflorescences (only known in fruit) rather few-flowered, c. 1.5-3 cm long; 
bracts minute, very early caducous; flowers 3 or 4(or 5)-valved, perianth glabrous, 
pedicel glabrous, at base not or only in a few cases indistinctly articulate. Male 
perianth globose or slightly depressed-globose, 1.5-2.0 x 2-2.5 mm, top and base 
(broadly) rounded, glabrous; pedicels 2-4 mm long, slender, glabrous; perianth at 
anthesis cleft to 43-2, not or but slightly collapsing on drying: valves 0.1-0.2 mm 
thick. Androecium subglobose or depressed-globose, 0.8-1.0 x 1.0-1.2 mm, top 
broadly rounded, circular in transverse section; anthers 7-9, almost completely 
sessile, incurved at the apex; central column broad and solid without an apical 
cavity Or minute, or rarely (see note) rather broad and shallow, 0-0.4 mm deep, 
sometimes with a short protuberance; androphore rather slender and distinct, c. 
0.3-0.4 mm long. Female perianth (only known from very young fruit) c. 3 mm 
long, 3-or 4 valved, glabrous; ovary glabrous. Fruits 2-4 per infructescence, ellip- 
soid, top (narrowly) rounded, base rounded, 3.0-5.5 x 2.5-4.5 cm, glabrous, drying 
blackish, finely granulate and sometimes + tuberculate, pericarp (4-)5-15(-20) mm 
thick; perianth not persisting under the fruit; stalk 5-7 mm long. 


Distribution. Peninsular Thailand, Malaya, Singapore, Sumatra. 

THAILAND. Peninsular: (Phusomsaeng 411) Fl. Thailand 40961. 

MALAYA (Perak, Kelantan, Selangor, Trengganu, Pahang, Negri Sembilan, Johore): FRI 0609, 
3530, 7819, 7982, 8346, 8909, 14412, 14463, 14500, 14750, 25024; KEP 76635, 80805, 94284, 94991, 
108878, 110378, 110406; Shah & Noor MS. 1760; SFN (Sinclair) 4015S. 

SINGAPORE. Ridley s.n.; Sinclair sen. (L) 


SUMATRA. Riau: Soepadmo 186; Jambi: Roos & Franken 1936. 


i Nitti ae 


New account of Horsfieldia 3 195 


Ecology. Primary and secondary dryland forest, also in periodically inundated 
forest; mainly on sandy soil, sandy loam soil, sandstone; 0-350 m alt. Flowers in 
March, August; fruits mainly May to July. 


NOTES 


1. Fieldnotes. Erect tree with clear bole, bole once recorded as + tapering, once 
recorded to have buttresses to | ft. Bark brown to blackish-brown, shallowly 
fissured or usually thinly flaking. Inner bark fibrous, pale reddish; copious watery 
or sticky pink-red juice; slash wood pale, cream to pale pink-brown, red-flecked. 
Fruits recorded as large, up to 11 cm with thick fleshy pericarp; seeds relatively 
small, up to 5 cm long; fresh pericarp yellow to orange red, often flushed pink, 
smooth, waxy, shining. 


2. Sinclair recognized the present species as a distinct species under the name H. 
sucosa. Unfortunately, the lectotype-specimens, chosen by Sinclair, belong to the 
species treated by Sinclair under the name H. bracteosa, the latter thus being a later 
synonym of H. sucosa. The other specimens of King’s syntype of H. sucosa belong 
to yet other species, as pointed out by Sinclair (1958, p. 418). Our present species, 
hence, has to be given a new name, with a new type-specimen. 


3. Dry fruits reach c. 5.5 cm; according to the field labels the fruits may reach up 
to 11 cm, with thick fleshy-juicy pericarps, which obviously shrink a lot on drying. 


4. H. sparsa is well characterized from resembling species like H. sucosa and H. 
pallidicaula by its stout whitish twigs, dispersed leaves with a rather chartaceous 
consistency, the leaves sometimes being crowded towards the thickish and short 
terminal leaf bud, the relatively long petioles; the inflorescences are nearly glab- 
rous, the flowers 3- or 4-merous, the pedicels at base not or only indistinctly 
articulate, the androecium is rather distinctly stalked (androphore) and generally 
without an apical cavity. Only in Phusomsaeng 411, from Peninsular Thailand, is 
the apical cavity in the androecium rather large, reminiscent of H. sucosa. The 
fruits are large, with a particularly thick and fleshy pericarp; the perianth is not 
persistent. H. sucosa has articulate pedicels; H. pallidicaula differs with its sessile 
-androecium. 


5. The epithet ‘sparsa’ alludes to the dispersed leaves, an uncommon feature in 
Horsfieldia. 


51. Horsfieldia triandra de Wilde, sp. nov. Fig. 1B(51); 23 


Folia elliptica ad oblonga, 5-9 cm longa. Flores masculi pubescentes. perianthio crasse-carnoso, 3- 


valvato, in anthesi usque ad c. % diviso, androecio + turbinato, c. 1.5 mm longo, antheris 3, pro 
majore parte sessilibus, c. 0.6 mm longis. — Type: Sumatra, Forbes 2465 (L). 


Tree-height unknown. Twigs terete or subterete, not or only faintly ridged, 
towards the top 1.5-3(-4) mm diam., dark brown, early glabrescent, tomentum 
rusty, with hairs c. 0.3-0.7 mm long, bark finely and lower down coarsely striate, 
not flaking, lenticels small but conspicuous. Leaves in 2 rows, membranous to 
thinly chartaceous, elliptic-oblong to oblong, broadest at or slightly below the 
middle, 5-9 x 2-3.5 cm, base (short-) attenuate, top acute-acuminate with conspi- 
cuous acumen 8-12 mm long; upper surface glabrous, drying dark olivaceous to 
dark brown, lower surface brown, glabrous (early glabrescent), without blackish 


~> 


ea ee 
em eS 
eR ah 
my r 


te nh png Se My OTIS ee swt 
BU ELE ee re a So ne 
Ps 


tune a teene ces 
Sr Sawer 


Fig. 23. Horsfieldia triandra de Wilde. 
a. habit of leafy twig with male inflorescences, note leaf-like bracts, x 2; b. mature male 
flower, X 6; c. ditto, longitudinally opened, showing thick-walled perianth, and androecium, 
x 6; d. androecium, longitudinal section, schematic, x 6. — a-d from Forbes 2465. 


dots; midrib raised above, glabrous; nerves 6-10 pairs, above flat or + sunken; 
tertiary venation forming a lax network not or scarcely visible on both surfaces; 
petioles 7-13 x 1.0-1.5 mm, early glabrescent; leaf bud c. 6-8 x 2 mm, densely rusty 
pubescent with hairs 0.3-0.7 mm long. Inflorescences densely rusty pubescent with 
hairs c. 0.3-0.4 mm, in C’: 1 or 2 times ramified; flowers not many (10-20 flowers 
per inflorescence and in different stages of development) 3-5 x 1-2 cm, common 
peduncle 3-10 mm long, flowers solitary or in loose clusters of 2-4 each; bracts 
ellipsoid-oblong, pubescent, 2-3 mm long, caducous, and usually with one (rarely 2) 
persistent subterminal bract, enlarged and resembling a small foliage leaf, 5-12 mm 
long (see notes). Flowers 3-valved, perianth towards the apex glabrous, lower 
down finely pubescent, pedicel pubescent with hairs 0.2-0.4 mm, inarticulate at 
base. Male perianth + obconical-obovoid, the top broadly rounded or with a blunt 
tip, base tapering, c. 2.5 x 2.5 mm, glabrous in the upper half, pubescent with hairs 
0.2-0.4 mm long towards the base; pedicel 3-4 mm long, slender; perianth at 
anthesis cleft for only c. %-%, not collapsing on drying, valves at apex c. 0.2, 
towards the base c. 0.4 mm thick, + clasping the anthers, at anthesis hardly 


196 


New account of Horsfieldia 3 197 


opening, the basal part of perianth thick-walled, 0.8-1.0 mm thick. Androecium 
incl. androphore + turbinate, 1.5-1.7 x 0.6-0.8 mm, subcircular in transverse 
section; anthers 3 (6 thecae), acutish, suberect, subsessile, 0.6-0.7 mm long, the 
apical c. 0.3 mm mutually free; androphore + obconical, tapering, c. 1.0 xX 
0.6-0.7(-0.8) mm, above continuing into the column. Female flowers and fruits not 
seen. 


Distribution. C. and S. Sumatra. 


SUMATRA. Central (West Coast): b.b. 6479 — South (exact locality not known): Forbes 2465. 
Ecology. Not known; the sterile collection b.b. 6479 collected at c. 1000 m alt. 


NOTES 


I. Forbes 2465, from Sumatra and without field data, is the only one which is 
fertile. Sinclair identified it at first as Pygeum (now Prunus), others as possibly 
Platea and finally as a Horsfieldia species. 


The mature male perianths deviate from most Horsfieldias by the obconical 
shape and the thick-leathery texture, the perianth opening only slightly at the very 
top, and the turbinate androecium with only 3 anthers at the top, which are clasped 
by the perianth-valves before anthesis. 


These flowers look as if they might be diseased, but on opening I found flowers 
and androecia normal. The rather few-flowered inflorescences, which bear one or 
two leaf-like enlarged bracts in the apical portion also look aberrant, but no trace of 
disease can be found. Enlarged leaf-like bracts in the inflorescences are occasional- 
ly found in the inflorescences of other species, e.g., in H. irya. 


2. H. sterilis from Borneo, H. pulverulenta from new Guina, H. crux-melitensis 
(to a lesser extent) and related species from new Guinea have a 2-valved perianth 
and the following combination of characters, i.e., flowers with a perianth equally 
thick-leathery, similarly the opening limited to the top of the perianth, and the 
androecium having a “reduced number”’ of stamens. 


3. A second specimen, b.b. 6279, from W. Central Sumatra, at c. 1000 m, is 
sterile. Sinclair determined it as H. ridleyana. 


52. Horsfieldia tristis de Wilde, sp. nov. Fig. 1B(52) 


Folia membranacea ad chartacea, in sicco tristia, gemmis tomento minus quam 0.1 mm longis. Cortex 
ramulorum pallide brunneus, striati, + delapsus. Perianthium masculum late ellipsoideum, c. 3 mm 
longum, 3-valvatum, in anthesi usque ad 5-4 diviso; androecio longiore quam latiore, antheris 12-20. 
— Type: Sarawak, Lai Shak Teck S. 37470 (L; iso: K; KEP, MO, SAN, S, n.v.). 


Tree 10-15 m. Twigs terete, not ridged, towards the top 3-5(-7) mm diam., early 
glabrescent, tomentum greyish to rusty, with hairs c. 0.1 mm long or less, bark 
lower down rather bright brown to yellowish, coarsely striate with a tendency of 
cracking longitudinally or flaking; lenticels smallish, conspicuous or not. Leaves in 
2 rows, membranous to thinly chartaceous, elliptic-oblong to lanceolate, broadest 
at or slightly above the middle, or + parallel-sided, 14-32 x 4-8.5 cm, base 


198 Gard. Bull. Sing. 38(2) (1985) 


attenuate, tip acute-acuminate; above drying dull and to a finely wrinkled or 
granulate structure, dark olivaceous or olivaceous-brown, lower surface glabrous, 
olivaceous-brown, without blackish-brown dots; midrib flat or slightly raised 
above, glabrous; nerves 11-17 pairs, above flat or slightly raised or sunken, the 
marginal arches indistinct or invisible; tertiary venation forming a + lax network, 
inconspicuous or invisible; petioles 6-12 x 2.0-5 mm, glabrous; leaf bud slender, 
10-15 x 1.5-2.5 mm, densely greyish to dull-brown pubescent with hairs up toc. 0.1 
mm long. Inflorescences glabrous or with a few scattered minute hairs up to c. 0.1 
mm, in CO: c. 3 times ramified, rather many-flowered, 5-12 X 3.5-7 cm, common 
peduncle 7-25 mm long; @ inflorescences (in fruit): c. 2-3 cm long, once or twice 
ramified. Flowers (C’) in loose clusters 2-5 each, 3- (or 4-)valved, perianth glabrous; 
pedicel glabrous, at base not articulate; bracts not seen, caducous. Male perianth 
obovoid-ellipsoid to broadly ellipsoid, 2.8-3.7 « 2.0-3.0 mm, top broadly rounded, 
based rounded to short-tapering; pedicel 1.5-5 mm long, slender and well marked 
off from the perianth; perianth at anthesis cleft to “5-4 (to nearly 3); valves 
0.2-0.4 mm thick. Androecium obovoid-ellipsoid, top broadly rounded or + de- 
pressed, in transverse section subcircular or bluntly 3- or 4-angular, 2.0-2.7 x 
1.5-2.0 mm; anthers 12-20, mutually appressed, almost completely sessile with free 
apices O0-0.1(-0.2) mm long, column broad and solid with a rather broad apical 
cavity, 0.5-0.8 mm deep, sometimes with a broad and flat base, almost completely 
concealed by the overcurved anthers; androphore narrow, 0.2-0.3 mm long, largely 
hidden by the anther-bases. Female flowers not seen. Fruits 4-8 per infructescence, 
ellipsoid, top subacute, base rounded, c. 1.5 X 1.2 cm, glabrous, drying blackish, 
without lenticels or tubercles, pericarp c. 1.5 mm thick; stalk 1-2 mm long; perianth 
not persisting. 


Distribution. Sumatra (E. Coast of Tapanuli), Lingga Arch. (Singkep Isl.), 
Borneo (Sarawak, S. Kalimantan). 


SUMATRA: Rahmat si Toroes 5486 — Lingga (Sinkep Isl.): Biinnemeyer 7100. 


BORNEO. Sarawak (lst. Div.): (Haviland) Kalong 1949; (Lai Shak Teck) S 37470 — §. Kalimantan 
(Sampit): Kostermans 8043. 


Ecology. Forest on flat land; 0-100 m alt. Flowers in August and November, 
fruits in September. 


Vernacular names. Kajoe penara (Singkep Isl.); Kajoe darodong lomba (Tapa- 
null). 


NOTES 
1. Fieldnotes. Flowers yellowish, fragrant. 


2. It seems closely related to H. fulva and allies of that on account of the similar, 
elongate, male flowers, but in H. fulva the leaves dry brown instead of olivaceous, 
the twigs brown, neither pale nor bright brown, the bark of the twigs does not tend 
to crack longitudinally, the inflorescences are pubescent, and the pedicels articu- 
late. 


3. The plants have rather yellowish twigs and pale, olivaceous leaves, and give a 
rather pale overall impression; twigs, leaves and inflorescences are almost com- 
pletely glabrous. Specimens from Borneo were identified by Sinclair as H. carnosa, 
the specimen from Singkep Isl. as H. glabra (see note 4), that from Sumatra as H. 
irya. 


New account of Horsfieldia 3 199 


4. The one from Singkep Isl., Bunnemeyer 7100, somewhat deviates by the 
relatively thin leaves, their upper surface not very distinctly dull and wrinkled and 
by the rather broadly ellipsoid, almost subglobose, male perianths; its androecium 
has a very broad apical cavity with a broad, almost flat base. 


53. Horsfieldia fulva (King) Warb. Fig. 1B(53) 


Myristica fulva King. Ann. Roy. Bot. Gard. Calc. 3 (1891) 297, pl. 124 — Horsfieldia fulva (King) 
Warb.,. Mon. Mynist. (1897) 297: Sinclair, Gard. Bull. Sing. 16 (1958) 396, fig. 37; 28 (1975) 33. — 
Syntype: Maingay 1304 (2426) (CAL, n.v.; iso: K), Scortechini 184 a (CAL, n.v.; iso: BM, K. L: G. 
n.v.). 


Tree 10-20 m. Twigs terete, towards the apex 3-5(-10) mm diam., rather late- 
glabrescent, tomentum dense, of short-dendroid hairs 0.2-0.3 mm long, lower down 
with the bark grey or grey-brown, finely to hardly striate; lenticels usually many, not 
very conspicuous; bark on older twigs not flaking. Leaves in 2 rows, chartaceous to 
coriaceous, dull, elliptic-oblong to (obovate-) oblong, broadest at or somewhat 
above the middle, 13-21 x 4-9.5 cm, base attenuate, top acute-acuminate to 
bluntish; upper surface drying dull pale olivaceous to bright brown, glabrous, lower 
surface pale brown, glabrous or with some persistent tomentum of hairs 0.2-0.3 mm 
on and near the midrib and lateral nerves; without larger brown-blackish dots; 
midrib above flattish or somewhat raised, early glabrescent; nerves 11-14(-18) 
pairs, above flat to + sunken, the lateral arches regularly shaped, not distinct: 
tertiary venation not or hardly visible on both surfaces; petioles 8-13 x 2.5-3.5 mm, 
glabrous; leaf bud 10-15 x 2.5-3 mm, densely pubescent with hairs c. 0.2-0.3 mm. 
Inflorescences situated generally behind the leaves, not very densely pubescent, the 
hairs stellate-dendroid, 0.2-0.3 mm, in CG: c. 3 times ramified, rather many- 
flowered, 3-10 x 2-6 cm, common peduncle 2-7 mm long; in 2: few-flowered, 1-2 
cm long; bracts not seen, caducous. Flowers solitary or in loose clusters of up to 5 
each, 3-valved, perianth and pedicel glabrous, the pedicels + articulated at base. 
Male perianth ellipsoid or ellipsoid-obovoid, 3.0-4.0 x 2.0-2.5 mm, top rounded, 
base + attenuate to rounded: pedicel 1-3 mm long, slender; perianth at anthesis 
cleft to c. 45-44; valves 0.2-0.3 mm thick. Androecium elongate-ellipsoid, above 
truncate to rounded, sub-cylindrical to + 3-angular, 2.0-3.0 x 1.1-3 mm: anthers 
10-12, almost entirely sessile, appressed, apices free up to 0.1 mm, the apex of the 
column narrowly hollowed or cleft to c. 0.3 mm, androphore rather narrow, up to 
0.1 mm long. Female perianth (according to Sinclair, 1959, p. 396): long-ellipsoid, 
5-6 mm long, 3-valved; pedicel stout, c. 3 mm long; ovary 2.5.-3 mm long, glabrous, 
stigma sessile, bi-lobed. Fruits up to 3 per infructescence, ovoid-ellipsoid, top 
obtuse to acutish, base broadly rounded, 2.2-2.4(-3.0) x 1.6-2.0(-2.5) cm, glab- 
rous, drying bright brown, without lenticels or warts, dry valves c. 3 mm thick; stalk 
c. 3-5 mm long; perianth persisting under fruit (always?). 


Distribution. Malaya (Perak, incl. Pulau Rumbia, Selangor, Negri Sembilan, 
Malacca), Sumatra (Jambi Prov.). 


MALAYA: FRI 16028, 16116, 25623, Kep. FN. 76031, 80627, 98837, 99334; Maingay (1304). 2426; 
Scortechini 184a; (Sinclair) SING 40170. 


SUMATRA: Jambi Prov. Roos & Franken T.F.B. 1510, 1526 


Ecology. Lowland rain forest: undulating country, on ridges; 0-200 m. Flowers 
and fruits throughout the year. 


200 Gard. Bull. Sing. 38(2) (1985) 


NOTES 


1. Fieldnotes. Bark yellowish-brown, thin, shallowly fissured longitudinally but 
not flaking; inner bark orange; wood white; sap watery, pale pink, not copious. 
Flowers orange, fruits yellow. 


2. Easily recognized by the dull parchment-like leaves when dry; the nerves 
above are flat or somewhat sunken, and the tertiary venation is not or hardly 
visible; it is one of the few species with a 3-valved perianth which is elongate and 
rather large, 3 mm long or more. I agree with Sinclair that it must be closely related 
to H. superba, which is larger in size in almost all aspects, and which has a generally 
persistent pubescence on the lower leaf surface. 


54. Horsfieldia superba (Hook. f. & Th.) Warb .....................0cc cena ee Fig. 1B(54) 


Myristica superba Hook. f. & Th., Fl. Ind. (1855) 162; A. DC., Prod. 14, 1 (1856) 194; Miq., Fl. Ind. 
Bat. 1 (2), 1 (1858) 62; Hook. f., Fl. Brit. Ind. 5 (1886) 105; King, Ann. Roy. Bot. Gard. Calc. 3 
(1891) 298, pl. 124 bis, 125 bis. — Horsfieldia superba (Hook. f. & Th.) Warb., Mon Myrist. (1897) 
295; Corner, Wayside Trees Mal. 1 (ed. 1940 & 1952) 476; Sinclair, Gard. Bull. Sing. 16 (1958) 393, 
fig. 36, pl. X B; 28 (1975) 141. — Type: Phillips s.n. (K; iso: LE. n.v.). 


Tree 10-30 m. Twigs terete or sometimes faintly angular, stout, towards the apex 
5-8(-13) mm diam., rather late-glabrescent, tomentum dense rusty, composed of 
dendroid hairs 0.5-1.0 mm long, lower down with the bark dark grey, finely striate, 
lenticels usually many and conspicuous; bark on older twigs sometimes slightly 
cracking and flaking. Leaves usually in 2, sometimes in 3 rows (see notes), 
coriaceous, dull, elliptic-oblong to oblong, broadest usually at about the middle 
(17-)25-40(-70) x (7.5-)10-18(-22) cm, base narrowly subcordate to short- 
attenuate, top bluntish to acute-acuminate; upper surface drying dull with finely 
wrinkled structures, olivaceous to grey-brown, glabrous or glabrescent (except 
sometimes the midrib), lower surface with brown or bright brown rather sparse to 
dense ‘“‘mealy” tomentum of dendroid hairs of mixed sizes, c. (0.3-)0.5-1.0 mm 
long, usually with many or few emerged hairs to c. 1.5 mm long; no larger 
brown-blackish dots present; midrib rather broad, flat above, pubescent or glabres- 
cent; nerves 15-25(-30) pairs, above flat or sunken, the lateral arches fairly regular- 
ly shaped, not very distinct above; tertiary venation not or hardly visible on both 
surfaces, petiole 6-15 x 5-7 mm, rather late glabrescent; leaf bud 20-30 x 5-10 mm, 
densely pubescent by hairs 0.5-1.5 mm long. Inflorescences behind the leaves, 
moderately densely + woolly pubescent with rather long-branched yellowish- 
brown dendroid hairs c. 0.5-1.0 mm long; in CG’: rather many-flowered, 2 or 3 times 
ramified, 7-15 x 2.5-10 cm, common peduncle 10-15 mm; in 9: + few-flowered, 
little branched, 2-5 cm long; bracts broadly ellipsoid, subacute-acuminate, up to 12 
x 10 mm, densely pale brownish pubescent, caducous. Flowers (0) up toc. 5 ina 
cluster, perianth 3- or 4-valved, glabrous, drying often with a grey-bluish tinge, the 
pedicels slender, glabrous, at base inarticulate. Male perianth ellipsoid to obovoid- 
ellipsoid, 6-7(-8) x 4-5 mm, top rounded, base rounded to short-attenuate, glab- 
rous; pedicel 1.5-3.5(-5) mm; perianth at anthesis cleft to c. 4-4; valves c 0.4 mm 
thick. Androecium elongate-ellipsoid, subtruncate above, the apex sterile (i.e., not 
bearing anthers) for c. 0.5 mm (always?), with a shallow tri-radiate crack, base 
subtruncate, subcylindrical to 3-angular in transverse section, c. (4.0-) 4.5-5 x 
2.2-2.5 mm; anthers 16-20, completely sessile, mutually appressed; androphore 
rather narrow, 0.1-0.4 mm long. Female perianth ellipsoid, c. 7.0-8.0 x 4.5 mm, 
glabrous, cleft at anthesis to c. 5, valves rather coriaceous, c. 0.5 mm thick; pedicel 
stout, 2-3.5 mm long, glabrous; ovary ovoid, slightly laterally compressed, c. 4-4.5 


New account of Horsfieldia 3 201 


x 3.0 mm, glabrous, stigma shallowly 2-lobed, broad, c. 0.5 X 1.5 mm. Fruits 1-3 
per infructescence, broadly ovoid-ellipsoid, top and base rounded, c. 3.8-5.5 xX 
2.8-4.5 cm, glabrous, drying dark brown, often + coarsely warty and wrinkled, 
valves + fleshy, when dry 8-12 mm thick; stalk stout, c. 3-6 mm long; perianth-parts 
persistent under the fruit. 


Distribution. Malaya (all states except three), Singapore, Sumatra (Central- 
West). 


MALAYA. Ahmat F.M.S. 4867; Curtis 2966; FRI 0739, 4511, 5416, 10512, 16040, 17076, 25695; 
Hervey s.n; Holmberg 2100; Kadim & Noor 364; Kep FN 70473, 80618, 93151, 94998, 98192, 109000; 
King’s Coll. 8024; Phillips s.n.; Ridley 10526; Scortechini s.n.; Shah & Noor MS 1894; SFN. 28703, 
40496, 40570; Soepadmo 758; Zainudin & Kasim 01687. 


SINGAPORE. SFN (SING) 36141, 40047, 40174, 40688. 


SUMATRA. b.b. 23701; Koorders 10383. 


Ecology. Forest on alluvial soils, undulating country, also in swampy forest; 
0-400 m alt. Flowers visited by bees (Whitmore FRI 4511). Flowers August to 
October, most fruits collected May to August. 


NOTES 


1. Fieldnotes. Bole straight. Bark longitudinally fissured or distantly dippled or 
cracked. Bark slash brittle, gritty; slash wood soft, white to yellowish. Leaves 
glossy above, becoming dull with drying. Flowers bright yellow, smell unpleasant 
or of ripe pears, visited by bees. Fruits globose, greenish-yellow, yellow, or orange. 


2. Sinclair suggested (l.c., p. 141) to place the present species together with the 
much related H. fulva in a separate series of their own, mainly on account of the 
oblong staminal column and the oblong or obovoid male perianth. 


3. I have not seen the collection Koorders 10383, in BO, from Central Sumatra. 
The specimen b.b. 23701, from Sumatra, West Coast, at 400 m, is sterile, and has 
the leaves in 3 rows. Apparently this is the top of a robust erect-growing orthrotro- 
pic sterile shoot. However, in Whitmore FRI 4511 (Malaya), fertile male-flowering, 
the leaves are — possibly by way of exception — arranged in 3 rows, and evidently 
not distichous as in the remainder of the material. 


4. Sterile specimens of Gymnacranthera bancana, also with a stout habit and 
tomentum remaining on the twig apex and the lower leaf surface, may be confused 
with the present species. However, in the former the hairs of the tomentum are 
much more interwoven forming a thin felty mat, whereas in H. superba the leat 
undersurface is covered with harsh, stellate-dendroid hairs. 


5. The fruits are recorded by Sinclair (1958, p. 395) as being as large as 7-9 X 
5.5-6.8 cm, and at first covered with harsh rusty scurf, becoming glabrous. In the 
material at my disposal the fruits are all smaller and glabrous, consistent with the 
glabrous ovary, and I assume that Sinclair’s description was probably based on a 
mixture of species. 


55. Horsfieldia sessilifolia de Wilde, sp. nov 


Horsfieldia species H. sylvestris similis, ab ea differt folius maioribus, c. 50 x 14 cm, dense breviter 


202 Gard. Bull. Sing. 38(2) (1985) 


tomentosis ad folia paginam inferiorem, petiolo brevissimo ac lato, usque ad c. 3 mm longo, 
perianthio 3-valvato, usque ad  diviso, ovario subglabro. — Type: Turkey bin Tran S 27808 (L; iso: 
K, SING, Silvic. Sibu, n.v.). 


Tree c. 30 m. Twigs terete, stout, towards the apex c. 8-10(-20) mm diam.; twig 
apex (and leaf bud) not seen; bark lower down glabrous, finely striate and rather 
densely set with conspicuous lenticels. Leaves arranged in 2 rows, chartaceous, 
oblong-lanceolate, + parallel-sided, c. 50 x 14 cm, base broadly rounded to 
subcordate, top acute-acuminate; upper surface drying dark brown, glabrous, 
lower surface finely bright-brown pubescent with densely interwoven hairs c. 
0.1(-0.3) mm; without dark brown dots; midrib above flattish; nerves c. 30(-35) 
pairs, above flat or slightly sunken, beneath with distinct submarginal arches; 
tertiary venation forming a coarse network, indistinct, on the lower leaf surface and 
largely hidden by the tomentum; petiole short, c. 2-3 X 8 mm, or leaves sessile; leaf 
bud not seen. Inflorescences only known in Q: c. 3 times ramified, c. 4-5 X 3.5 cm, 
rather many-flowered; branches rather densely rusty woolly-pubescent with hairs c. 
0.3-0.5 mm; common peduncle c. | X 0.8 cm; bracts oblong-lanceolate, c. 1 cm 
long, inside subglabrous, outside pubescent as the inflorescence, caducous. Flowers 
in Q in clusters of 2-5 each; perianths 3-valved, minutely pubescent in the lower 
half. Male perianth not seen. Female perianth obovoid-ellipsoid, stout, c. 4.5-5.0 x 
4.0-4.5 mm, coriaceous, pubescent with hairs c. 0.1 mm in the lower half, cleft at 
anthesis to c. ¥s-Y10 only, valves c. 0.5 mm thick; pedicel stout, c. 1 mm long, 
minutely pubescent; ovary ovoid, c. 2.5 X 2.5 mm, subglabrous, with a few minute 
hairs, only on and near the base of the suture, whitish, less than 0.1 mm long; 
stigma broadly 2-lipped, c. 0.2-0.3 x 1.5 mm, not or only faintly lobed. Fruits not 
seen. 


Distribution. Borneo. Sarawak: Sibu district (3rd Div.), known only from the 
type from Sg. Tutus, Loba Kabang (S) P.F., Batang Igan. 


Ecology. Lowland mixed swamp forest, apparently under 100 m alt. Flowers in 
June. 


Vernacular name. Kumpang tembaga. 


NOTES 
1. Fieldnotes. Recorded as a tree 100 ft. tall, 42 ins. girth, with stilt roots. 


2. The specimen on which the new species is based is reminiscent of some other 
stout-leaved Horsfieldias, viz. H. sylvestris from SE. Malesia, and to a lesser extent 
of H. splendida (Borneo), H. superba (Malaya, Sumatra), or H. pulcherrima 
(Sumatra), but the present new species is distinct in its almost sessile leaves and the 
densely short-pubescent lower leaf surface. H. sylvestris is generally less stout; it 
has similarly subsessile (but narrower) leaves and 2-valved perianths. The other 
mentioned stout-leaved species all have the leaves distinctly petioled. 


3. The specimen was collected in June, 1971, i.e., after Sinclair’s work was 
published. In the absence of male flowers, Mr. J. Koster (Koster & Baas, 1981) 
investigated the leaf anatomy and confirmed the identity of the specimen as being a 
Horsfieldia. 


New account of Horsfieldia 3 203 


56. Horsfieldia grandis (Hook. f.) Warb. Fig. 1C(56) 


Myristica grandis Hook. f., Trans. Linn. Soc. 23 (1860) 157 — Horsfieldia grandis (Hook. f.) Warb., 
Mon. Myrist. (1897) 301; Sinclair, Gard. Bull. Sing. 16 (1958) 400, f. 39; 28 (1975) 48. — Type: 
Sabah, Low s.n. (K). 


Myristica rubiginosa King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 302, pl. 130 — Type: Singapore, King’s 
Goll 1233 (CAL, nwi31s0:-K;.L), 


Tree 6-25 m. Twigs stoutish, terete, towards the top (3-)4-10 mm diam., late- 
glabrescent, tomentum dense, harsh, with hairs c. 1.0-1.5 mm long; bark of twigs 
lower down rather coarsely striate, when older longitudinally and sometimes slight- 
ly transversely cracking, hence slightly flaking; lenticels often present, somewhat 
elongate, not conspicuous. Leaves in 2 rows, membranous or chartaceous, some- 
what bullate, elliptic-oblong to oblong-oblanceolate, broadest usually somewhat 
above the middle, 12-40 x 5.20 cm, base + attenuate to subcordate, top acute or 
acute-acuminate; upper surface drying olivaceous to dark brown, minutely paler 
pustulate, largely with harsh, persistent tomentum of rather distant hairs, some- 
times glabrescent, but always scabrous from harsh, persistent hair-bases, lower 
surface with persistent tomentum of dense to rather distant hairs c. 1.0-2.0 mm 
long, harsh-woolly to the touch; without brown dots; midrib above usually densely 
pubescent, somewhat raised; nerves (8-)10-16(-19) pairs, above flattish or usually 
sunk, the marginal arches distinct; tertiary venation forming a lax network, well 
visible on both surfaces especially the lower; petiole 6-15 x 2.5-6 mm, densely 
pubescent; leaf bud short-conical, densely pubescent, 7-15 mm long. Inflorescences 
densely pubescent, hairs yellow-rusty, 1.5-3.0 mm long, in C’: many-flowered, 3 or 
4 times ramified, usually rather lax, c. 6-25 X 2.5-10(-15) cm, common peduncle up 
to 12 mm long; 9 inflorescences 1.5-5 cm long, the flowers often of different age 
and size; bracts generally oblong to lanceolate-linear, (1.0) 3-12 mm long, caduous. 
Flowers 3- or 4-valved, glabrous, solitary or usually in loose clusters, often aggre- 
gated into compound clusters corresponding to the main ramifications of the 
inflorescences; pedicels slender, glabrous, at base inarticulate. Male perianth glo- 
bose or depressed-globose, 1.2-1.8(-2.0) mm diam, at apex rounded or broadly 
rounded, base rounded; pedicel (0.5-)1-2 mm long; perianth at anthesis cleft to c. 
’2 to nearly 2-way, valves c. 0.2 mm thick. Androecium depressed globose, not 
laterally compressed, apex broadly rounded to depressed, c. 0.5-1.0 x 0.8-1.5 mm; 
anthers 8-10, almost completely sessile, incurved towards the top, column broad, 
solid, hollow c. 0.2 mm deep; androphore rather narrow, 0.2-0.4 mm long. Female 
perianth subglobose to broadly ellipsoid, 2.0-3.2 x 2.0-2.8 mm, cleft at anthesis to 
c. 44-3, valves 0.2-0.3 mm thick; pedicel 0.3-0.5 mm long, glabrous except some- 
times with a few hairs at the base; ovary globose to broadly ellipsoid, c. 1.5-2.0 x 
1.5-1.8 mm, glabrous, stigma sessile, faintly 2-lobed, c. 0.2 mm high. Fruits 2-10 
per infructescence, + clustered, obovoid-ellipsoid, top rounded 1.0-1.4 x 0.8-1.1 
cm, glabrous, valves c. 1.5 mm thick, drying dark brown or reddish brown, without 
distinct lenticels, not warted; seed almost globose, 8-10 mm diam.; stalk c. 1 mm 
long; perianth persistent. 


Distribution. Sumatra (Palembang; Riouw, n.v.), Malaya (Pahang, Johore), 
Singapore, Borneo (Sarawak, Brunei, Sabah, E. Kalimantan). 


MALAYA. Pahang: Chew Wee-lek & Noor, CWL 261 — Johore: FRI 8699, 17135; Holttum 9304; 
Ridley 4827; SFN 36831; Shah, Noor & Shukor MS 2060; Sinclair 10602. 


SINGAPORE. King’s Coll. 1233; Ridley 4133; Sinclair 9363. 


SUMATRA. Palembang: Endert 44 — Riouw Arch.: Teysmann s.n. (not seen). 


204 Gard. Bull. Sing. 38(2) (1985) 


BORNEO. Sarawak: Hose 655; S 15006, 18940, 23057, 25486, 26257, 28966, 29189, 37731, 37952 — 
Brunei: Ashton BRUN 64, 752, 3011; Fuchs & Muller 21160; Wyatt-Smith KFN. 80106: Sinclair & 
Kadim 10455 — Sabah: Castro A 83; SAN A 1741, 3691, 17497, 36338, 36914, 44361, 44713, 48857 
69297, 72199, 77512, 80921, 84636, 93684; Sinclair, Kadim, & Kapis 9249; Wood 9; Wood 1226 — E. 
Kalimantan: Endert 5081; Jaheri 727; Korthals 92; Kostermans 7035, 7044. 


Ecology. Primary and secondary forest, ridge forest; on sand and clay soil, 
sandstone; 0-600 m alt. Flowers and fruits throughout the year. 


Vernacular names. Kumpung (Malay, Iban name; Sarawak); Penarahan 
(Brunei); Tjemanding (Sumatra, Palembang). 


NOTES 


1. Fieldnotes. Slender tree, without buttresses; monopodial branches, branchlets 
few, horizontal. Bark sometimes recorded as smooth, non-flaking, usually as 
longitudinally fissured, or scaly, or flaky, or cracked; strips c. 1 mm thick, c. 1 cm 
wide, hard and thin; inner bark c. 2-3 mm thick, slash rich red-brown, with reddish 
watery exudate. Sapwood soft, whitish to yellowish pink, wood pale brown. Flow- 
ers yellow, with faint odour. Fruits yellow-green, yellow or + orange, pericarp 
inside pink; aril orange; seed pale grey. 


2. Perianths are usually 4-, less frequently 3-valved. I think there are 8-10 
anthers, but Sinclair says 13-15, the discrepancy is probably caused by the thecae 
being tightly appressed and difficult to count. 


3. The fruits are small, rather contrasting with the stout habit of twigs and 
leaves. The perianth remains persistent under the maturing fruit. 


4. The circumscription of the present species is the same as that by Sinclair. It 
belongs apparently to the group of West-Malesian species with pubescent lower 
leaf-surface, and is probably most closely related to H. flocculosa. H. grandis is 
well-marked in many details and readily recognizable by the scabrous upper leat 
surface which is caused by the harsh hair-bases. 


57. Horsfieldia wallichii (Hook. f. & Th.) Warb. Fig; 1CCST) 


Myristica wallichii Hook. f. & Th., Fl. Inc. (1855) 161 (p.p., Wall. Cat. 6806 being a mixture, see note 
by Sinclair, loc. cit., p. 158); A. DC., Prod. 14, 1 (1856) 230; Miq., Fl. Ind Bat, 1 (2), 1 (1858) 67; 
Hook. f., Fl. Br. Ind. 5 (1886) 105; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 303, pl. 132 & 133, 
excl. syn. M. crassifolia Hook. f. & Th. — M. horsfieldia auct. non BI1.: Wall. Cat. (1832) 6806, p.p. — 
Horsfieldia wallichii (Hook. f. & Th.) Warb., Mon. Myrist. (1897) 305; Sinclair, Gard. Bull. Sing. 16 
(1958) 405, fig. 41, pl, XI A; 28 (1975) 156 (p.p. excl., part of the Bornean material = H. borneensis) 
— Syntype: Malacca, Griffith s.n. (K); Singapore, Wallich Cat, 6806, p.p. (CAL n.v.; K); Lobb s.n. 
(K, n.v.). 


Tree, 10-30 m. Twigs terete or drying somewhat flattened, usually conspicuously 
hollow, towards apex 3-6(-9) mm diam., early to late-glabrescent from tomentum 
of hairs c. 0.3-0.6 mm, bark lower down usually dark brown or blackish, coarsely 
striate, with or without inconspicuous lenticels; bark on older twigs often longitudi- 
nally fissured, sometimes flaking. Leaves in 2 rows, membranous to coriaceous, 
ovate-oblong to oblong-lanceolate, broadest usually at about the middle or some- 
what parallel-sided, (14-)19-40 x (4-)4.5-12 cm, base rounded to short-attenuate, 


New account of Horsfieldia 3 205 


top subobtuse to acute-acuminate; upper surface drying olivaceous to blackish- 
brown, glabrous (with minute tomentum remaining on the midrib or not); lower 
surface late-glabrescent or tomentum locally persisting, of rather spaced to dense 
dendroid hairs (0.3-)0.5-0.8 mm long, always provided with typical scattered 
brown-black dots and stripes of irregular sizes; midrib above flat or slightly raised, 
late glabrescent: nerves (12-1) 15-28 pairs, above very slender, flat or sunken; 
tertiary venation forming a lax network, usually indistinct or invisible, petioles 
relatively long, 15-35 x 2.5-4.5 mm, glabrescent; leaf bud c. 20-30 x 4-6 mm, 
densely pubescent, hairs c. 0.3-0.6 mm. Inflorescences usually behind the leaves, 
moderately to densely pubescent with stellate-dendroid hairs 0.5-1.0 mm long, in 
CO: large, many-flowered, 3 or 4 times ramified, 10-33 x 6-22 cm, common 
peduncle 30-70 mm, in Q: stoutish, fewer-flowered, c. 3-7 cm long; bracts broadly 
ovate, 3-10 mm long, densely woolly-pubescent, caducous. Flowers in CO’ in clusters 
of 5-12, perianth 3-(or 4)-valved, glabrous, or in 9 glabrescent, pedicel short, 
pubescent, at base inarticulate. Male perianth broadly obovoid, 2-2.5(-3.0) x 
2.4-2.5 (-3.0) mm, top broadly rounded, base + attenuate, glabrous; pedicel short, 
0.3-0.6 (-1.0) mm, pubescent with hairs 0.2-0.3 mm long; perianth at anthesis cleft to 
c. 43-*4; valves c. 0.2-0.3 mm thick. Androecium broadly obovoid to subglobose, 
above + depressed with a tri-radiate crack, based rounded to attenuate, faintly 
3-angular in cross-section, 1.3-2.0 xX 1.5-2.0 mm; anthers (12?-)15-23, completely 
connate, sessile and closely appressed, at apex rather deeply incurved into the 
broad hollow c. 2-way deep in the column; androphore short and narrow, up to 0.3 
mm long. Female perianth ovoid-ellipsoid, 2.5-4.0 x 2.0-3.5 mm, glabrescent from 
hairs 0.1 mm, cleft at anthesis to c. ¥%, valves c. U.S mm thick; ovary ovoid to 
subglobose, 2.0 x 2.0-2.5 mm, glabrous, stigma rather broad, faintly 2-lobed, c. 0.3 
x 0.8 mm; pedicel stout, 0.5-1.5 mm long, densely pubescent by hairs 0.2-0.3 mm. 
Fruits 2-9 per infructescence, ovoid-ellipsoid, top and base rounded, 4.0-6.0 x 
3.0-4.5 cm, glabrous, drying dark brown, smooth or wrinkled, not or only faintly 
warted, valves 10-15 mm thick; stalk stout, 4-6 mm long; perianth generally 
persistent under the fruit. 


Distribution. Malaya (all provinces except Perlis and Negri Sembilan), Singa- 
pore, Sumatra (Aceh, Tapanuli, W. & E. Coast, Indragiri, Djambi, Palembang; 
Simalur (Simeulué), Morsala, Mentawai Isl., Bangka), Borneo. 


MALAYA. FRIO514, 2135, 4565, 6703, 13252, 14675, 14771, 17076; Kep. FN. 94671, 99010; Kunstler 
(King’s Coll.) 4827; Maingay 2665; Scortechini 246a; SFN 40484; Sinclair 9984. 


SINGAPORE. Maingay 1001, 1001A, 1283, 1284; SFN. 33556, 34439, 39486, 39487, 40216; Wallich 
Cat. 6806. 


SUMATRA. Achmad 251, 514, 675, 1219; b.b. T 944, 16382, 17452, 19346, 24049, 30033, 31758, 
32146; Forbes 3048, 3078; Kostermans 205 (b.b. 34139), b.b. 34063, b.b. 34194; Kostermans & Anta 795, 
845, 1068, 1194; Krukoff 4114; Mochtar 62A, 82A, 91A 97A; de Wilde & de Wilde-Duyfjes 18931. 


BORNEO. psarawak: Haviland 2182 — Sabah: BNB For. Dept. 1682 — Kalimantan, West: b.b. 
25505; E. and SE.: b.b. 16139, 21179; Forman 484; Kostermans 5651; Leighton 844; Hubert Winkler 
2419. 


Ecology. Lowland forest, on ‘red’ soil, granitic sand soil, loam soil with coral 
limestone; ridge-top forest; 0-470m. Flowers and fruits throughout the year. 


Vernacular names. Asem-asem (Bangka, Malay), Piangu, Pijangu (Bangka, 
Malay); Satim, Soemarallah-falah, Soemarallah-oeding, Toetoen soemarallah deé- 
lok (all Simeulué Isls.). 


206 Gard. Bull. Sing. 38(2) (1985) 


NOTES 


1. Fieldnotes. Bole straight; crown dense, monopodial. Bark shallowly or deeply 
longitudinally fissured, dark grey, not flaking. Bark c 1 cm. thick, slash under bark 
bright red, pink, reddish brown, or deep red; slash wood whitish, pale or dirty 
yellowish, light brown, brown or red-brown. Flower buds blue-green or yellow at 
anthesis. Fruits glaucous, turning green-yellow, yellow, orange, orange-yellow, or 
red; aril orange. 


2. This species is identical with Sinclair’s circumscription of the species for the 
Malay Peninsula (Sinclair, 1958), but if compared to Sinclair’s, 1975, the majority 
of the specimens from Borneo should be excluded. These specimens are presently 
described as a new species H. borneensis. 


3. H. wallichii is, in the sterile state, always recognizable by the blackish dots 
and stripes scattered on the lower leaf surface. Similar dots are only found in a few 
other species, incl. H. borneensis, but the latter differs in general habit and the 
male flowers; it has almost similar fruits. 


H. wallichii in the sterile and fruiting state can also be confused with H. superba, 
a large-leaved species, but this lacks the blackish markings on the lower leaf 
surface. H. motleyi has also somewhat similar leaves, 1.e., drying dull above, and 
with flat or sunken lateral nerves; apart from the difference in the flowers and 
fruits, its lower leaf surface is not dotted. 


58. Horsfieldia pulcherrima de Wilde, sp. nov. Fig. 1C(58) 


Ramuli robusti, tarde glabrescentes, pilis 1-1.5mm longis. Folia 24-36 cm longa, subtus pubescentia 
atque munita punctis sordide brunneis non-traumaticis. Perianthium masculum depresso-globosum, 
1-1.3 mm diam., 3-valvatum, glabrum, pedicello basi non-articulato. Androecium depresso- 
globosum, 0.8-1.0 mm diam., anthers 12. — Type: Malaya, Cockburn FRI 8008 (L; iso: K). 


Tree 7-27 m. Twigs stout, terete, towards apex 5-8(-14) mm diam., densely felty 
to woolly pubescent with rusty or reddish brown hairs c. 1.0-1.5 mm, late glabres- 
cent, bark coarsely striate, when older somewhat cracking and flaking; lenticels 
only in the older wood, large, not contrasting in colour. Leaves in 2 rows, char- 
taceous, elliptic-oblong to oblong, broadest usually at about the middle, (18-)24-36 
x 8-14 cm, base rounded to attenuate, top acute-acuminate; upper surface drying 
dull olivaceous brown to blackish brown, glabrous (glabrescent, and not scabrous), 
lower surface with dense rusty or red-brown tomentum of mostly dendroid hairs 
1.0-1.5 mm long and with scattered dark brown to blackish + wart-like marks (cork 
warts); midrib above rather slender, late-glabrescent, flat; nerves 18-23 pairs, 
above slender, flat to sunken, marginal arches not very distinct; tertiary venation 
forming a rather lax network, usually faint or scarcely visible; petiole 15-25 x 5-7 
mm, pubescent; leaf bud c. 25-30 x 7-9 mm, with hairs 1-1.5 mm long. Infloresc- 
ences behind the leaves, densely woolly-pubescent with + shaggy hairs 0.5-1.5 mm 
long, in &: 5-8 X 5 cm, many-flowered, 3 or 4 times ramified, common peduncle 
2-10 mm long; in 9 a short, irregularly shaped, woody knob (as in some Knemas), 
c. 1 cm long. Flowers subsolitary or in loose clusters of up to 15 each, 3-(or 
4-)valved, glabrous; pedicels glabrous, at base inarticulate; bracts ovate-oblong, 
acute, densely pubescent outside, 3-7 mm long, caduous. Male perianth somewhat 
depressed-globose, c. 1.0 x 1.2-1.3(-1.4) mm, top + depressed or broadly rounded, 
base rounded, pedicel 1-1.5 mm long, slender, not tapered; perianth at anthesis 


New account of Horsfieldia 3 207 


cleft to c. ¥% to nearly ¥2-way, valves c. 0.2 mm thick. Androecium depressed- 
globose, above + depressed and with a 3-(or 4-)radiate crack, 0.5-0.6 x 0.8-1.0 
mm; anthers 12 or 13 (c. 24 thecae), mutually closely pressed, completely sessile, 
the apical parts overarching the apical hollow to c. ¥2-way deep; column broadly 
saucer-shaped, androphore narrow, 0.1-0.2(-0.3) mm long, largely hidden by the 
anthers. Female perianth (as known from caducous remnants under fruit): c. 3 mm 
long, sparingly pubescent with hairs c. 0.3 mm long, at anthesis cleft to nearly 
Ya-way, valves c. 0.3 mm thick. Ovary pubescent. Fruits in clusters of 1-3 per 
infructescence, broadly ellipsoid to globose, c. 1.6-1.8 x 1.5-1.7 cm, densely shaggy 
rusty pubescent with hairs c. 2 mm long, valves + woody, c. 1.5 mm thick, seed 
broadly ellipsoid, c. 10 mm long, aril entire, completely enveloping the seed; stalk 
0-1 mm; perianth not persistent. 


Distribution. Malaya (Pahang, Johore), Sumatra (Jambi). 
MALAYA: FRI 8008, 19881, 31807; Kep. 104997, 110369. 


SUMATRA: Roos & Franken TFB 1983 (sterile). 


Ecology. Lowland primary forest, swamp forest; 50-600 m alt.; apparently a rare 
species. Flowers in March, fruits in June and September. 


NOTES 


1. Fieldnotes. Slender tree, crown monopodial. Bole straight; bark grey-brown 
to blackish, shallowly fissured, occasionally flaking. Inner bark red, laminated, 
with some red exudate. Slash wood whitish to pale yellow, wood brown. Immature 
flowers green. Fruits yellowish-brown hairy. 


2. All the specimens of this remarkable and beautiful new species known to me 
were collected after Sinclair’s revision of Horsfieldia. In sterile state the species 
may be confused with H. superba, H. flocculosa, or H. wallichii, but its CO flowers 
are quite different, rather resembling those of H. grandis. The species is peculiar 
because of its smallish, subglobose, densely tomentose fruits. In all related species 
the fruits (and ovary) are glabrous. 


3. If sterile, H. pulcherrina may superficially resemble and be confused with the 
stout-leaved Gymnacranthera bancana, but the latter has a different tomentum, 
and lacks the blackish dots on the lower leaf surface. 


59. Horsfieldia flocculosa (King) Warb. Fig. 1C(59) 


Myristica flocculosa King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 302, pl. 131 — Horsfieldia flocculosa 
(king) Warb., Mon. Myrist. (1897) 297; Sinclair, Gard. Bull. Sing. 16 (1958) 398, f. 38. — Type: 
Malaya, Selangor, King’s Coll. 8618 (CAL, n.v.: iso: BM, K, L, P; B+, Fl, G, KEP. Z, n.v.). 


Tree 10-28 m. Twigs stout, terete, towards the apex 6-10(-12) mm diam., densely 
felty-woolly pubescent with yellow brown or pale brown hairs (1.0-) 1.5-2 mm, late 
glabrescent, bark coarsely striate, soon longitudinally cracking, in older wood also 
transversely cracking and + flaking; lenticels absent or indistinct. Leaves in 2 rows, 
chartaceous, oblong to oblong-lanceolate, broadest at about the middle, 18-40(-45) 
x 6-13(-18) cm, base broadly to narrowly rounded or subcordate, tip acute- 
acuminate; upper surface drying usually olivaceous to (light) brown, finely pustu- 


208 Gard. Bull. Sing. 38(2) (1985) 


late and wrinkled, glabrous, not scabrous, lower surface with dense perisistent 
woolly tomentum of hairs 1.5-2 mm long, without blackish dots; midrib flat above; 
nerves 15-20 pairs, above sunken, marginal arches rather distinct and regularly 
shaped; tertiary venation forming a lax network, + faint above, on lower surface 
much obscured by the tomentum; petiole 7-14(-20) x 5-7 mm, densely pubescent; 
leaf bud stout, 10-15 mm long. Inflorescences behind the leaves, densely woolly- 
pubescent, in CO’: rather stout, many-flowered, 3 or 4 times ramified, (8-) 12-20 x 
5-14 cm, common peduncle stout, 3-5 mm diam., 1-10 mm long; in Q: little 
ramified, 1.5-3 cm long, rather few-flowered. Flowers solitary or 2-3(-4) together, 
generally not clustered, glabrous, (3- or) 4-valved in CO’, (2- or) 3-valved in Q; 
pedicels at base inarticulate, glabrous; bracts ovate to lanceolate, densely pubes- 
cent, 5-20 mm long, caducous. Male perianths broadly ellipsoid to obovoid, or 
sometimes subglobose, not or slightly laterally compressed, (2.0-) 2.2-3.0 X 2.0-2.7 
mm, top broadly rounded, base rounded, pedicel slender, not tapered, (1.5-)3.0- 
4.0 mm long; perianth at anthesis cleft to “%-%4, valves c. 0.2 mm thick. 
Androecium + ellipsoid or broadly obovoid, subtruncate, c. 1.2-1.5(-2.0) x 1.0- 
1.3(-1.5) mm, not or only little laterally compressed, base broadly rounded, often 
faintly 4-angular in transverse section; anthers 10-13 entirely sessile, (not? septate), 
the apices slightly curved into the shallow hollow, c. 0.2-0.3 mm deep in the apex of 
the broad and solid column; androphore short, c. 0.1 X 0.4 mm, usually hidden by 
the slightly sagged anthers. Female perianth broadly ellipsoid, c. 3.5 x 3.0 mm, 
cleft at anthesis to c. %-'5, valves c. 0.4-0.5 mm thick; pedicel stoutish, c. 2 mm 
long, glabrous; ovary broadly ovoid, c. 1.8 X 2.0 mm, glabrous except for a few 
minute hairs (always?) on the suture below the stigma, stigma minute, faintly 
2-lobed, c. 0.3 mm long. Fruits (almost mature, according to Sinclair) subglobose 
to slightly ellipsoid, glabrous, c. 3.0 x 2.5 cm, pericarp c. 5 mm thick; stalk 5 mm 
long; perianth (at first) persistent. 


Distribution. Malaya: Only seen from Selangor, Negri Sembilan, Pahang and 
Johore. 


MALAYA. (Whitmore) FRI 0048; Hassan & Kadim H. 98; Kep. 104653, 110225, 77746; King’s Coll. 
8618; SFN 16394, 32314. 


Ecology. Lowland rain forest, also swampy forest, regenerated forest; 0-300 m 
alt. Flowers mainly February-June. 


NOTES 


1. Fieldnotes. Buttresses absent. Bark distantly superficially fissured, blackish 
brown; inner bark pinkish brown, laminated; sapwood whitish. Exudate watery, 
red. Young leaves flocculose. Margins of leaves slightly revolute when dry. Leaves 
somewhat bullate, thickish, shiny medium green above, golden below. Flowers 
yellow, or waxy light yellow. The perianths are described by Sinclair (1958, p. 400) 
as ‘covered with circles which are hyaline in the centre and brown round the 
circumference’. 


2. Apparently much related to H. superba, H. fulva (the flowers + similarly 
shaped as the ©’), and also to the group of H. grandis, and others but is disting- 
uished by many characters. The species is presently accepted in the same sense as 
by Sinclair and earlier authors. 


New account of Horsfieldia 3 209 


60. Horsfieldia motleyi Warb. Fig. 1C(60) 


Horsfieldia motleyi Warb., Mon. Myrist. (1897) 304; Merr., En. Born. J. Str. Br. R. As. Soc. spec. 
number (1921) 268; Sinclair, Gard. Bull. Sing. 28 (1975) 81 — Myristica motleyi (Warb.) Boerl., 
Handl. Fl. Ned. Ind. (1900) 85. — Type: SE. Borneo, Banjermasin, Motley 355 (K, iso. CGE, n.v.). 


Horsfieldia macrobotrys Merr., Pl. Elm. Born. in Univ. Cal. Publ. Bot. 15 (1929) 76 — Type: Sabah, 
Tawau, Elmer 21882 (PNH, 7; iso: BM, K, L; BO, SING, and others not seen). 


Tree 12-35 m. Twigs terete, towards the apex (2.5-)3-5(-10) mm diam., late- 
glabrescent from dense rusty tomentum of hairs c. 0.5(-1.0) mm, bark lower down 
finely striate; lenticels not conspicuous; bark of older twigs not flaking. Leaves in 2 
rows, membranous, not bullate, elliptic to oblong, broadest usually at about the 
middle, 9-27 x 4-12 cm, base rounded to attenuate, top obtusish to acute- 
acuminate; upper surface drying dull dark-olivaceous to greenish brown, glabrous, 
lower surface with persistent rather dense tomentum of stellate-dendroid hairs of 
mixed size, c. 0.3-1.0 mm long, without larger blackish dots; midrib flattish above, 
sometimes late-glabrescent; nerves 9-21 pairs, slender above, often late- 
glabrescent, flat or usually sunken, the lateral arches not very distinct; tertiary 
venation forming a lax network, faint above; petioles 13-22 x 1.5-4 mm, rather 
late-glabrescent; leaf bud c. 15-20 x 3-4 mm, densely pubescent by hairs c. 0.5 mm 
long. Inflorescences densely pubescent with stellate-dendroid yellow-brown to 
rusty hairs c. 0.5-1.0 mm long, in C&’: 4 or 5 times ramified, many-flowered, 12-20 x 
10-14 cm, common peduncle 20-40 mm, the flowers before maturity often densely 
clustered and in submature buds often + angular; in 2: much ramified, rather 
many-flowered, 3-6(-10) cm long; bracts densely pubescent, broadly ellipsoid, 
acutish, 2-5 mm long. Flowers in clusters of 5-20, 3-valved, perianth and pedicel 
pubescent by hairs 0.1-0.3 mm, the pedicels slender, at base inarticulate. Male 
perianth subglobose or broadly obovoid, 0.8-1.0 x 0.7-1.0(-1.1) mm, top broadly 
rounded, base rounded, pubescent or late-glabrescent; pedicel slender, 1-1.5 mm 
long; perianth at anthesis cleft to c. half-way; valves c. 0.1 mm thick. Androecium 
(incl. androphore) broadly obovoid, above somewhat flattish and depressed in the 
centre, sub-circular in transverse section, 0.5-0.7 x (0.3-)0.5-0..6 mm; anthers 5, 
almost completely sessile, towards the top not or hardly incurved, 0.25-0.4 mm 
long; column broad, solid except for the shallow, broad, apical hollow to c. “10; 
androphore conspicuous, about as long as the anthers, broad, tapering, c. 0.2-0.3 
mm long, continuous with the anthers. Female perianth ellipsoid, c. 2.4 x 1.8-2.0 
mm, pubescent, cleft at anthesis to c.%4-%, valves c. 0.2 mm thick; pedicel c. 1.5 
mm long; ovary broadly ellipsoid, 1.5 x 1.2 mm, top and base broadly rounded, 
conspicuously grooved at one side, glabrous, stigma broad, shallowly 2-lobed, c. 
0.2 mm high. Fruits 5-15 per infructescence, broadly ellipsoid, top and base 
rounded, 1.9-2.3 x 1.6-1.8 cm, glabrous, drying brown, without lenticel-like wart- 
lets, dry valves c. 2 mm thick; stalk 2-4 mm long; perianth not persistent under 
mature fruit. 


Distribution. Whole of Borneo: Sarawak, Sabah, W., S., SE. and E. Kali- 
mantan. 


BORNEO. Sarawak: S (Chai) 36774, (Martin) 37945 — Sabah: Elmer 12882; San. 19007, 27443, 
29966, 77428 — Kalimantan, West: Hallier 347 — South: Kostermans 8117 — SE.: b.b. 18449, 26178; 
Korthals (39); Motley 355 — East: Kostermans 4864, 6859, 10428. 


Ecology. Primary forest, disturbed forest, poor forest, Dryobalanops forest, or 
hill Dipterocarp forest, usually on dry sandy soils, sandy clay, also loam with lime; 
often on ridge tops; 0-600 m alt. Flowers and fruits throughout the year. 


210 Gard. Bull. Sing. 38(2) (1985) 


NOTES 


1. Fieldnotes. Bole sometimes with steep buttresses to 5 m high, c. 30 cm out, c. 
9 cm thick, merging into the stem. Bark usually distinctly fissured, often scaly, 
strips 3-4 cm wide, to c. 10 mm thick, dark brown, black-brown, chocolate, or 
red-brown; living bark 10-12 mm thick, undulate in cross section, dark brown, or 
brown-red, inner bark red-laminated; cambium pinkish. Sapwood whitish, reddish 
white, heartwood reddish or pinkish or brown; exudate of bark a red watery 
latex, sometimes recorded as sticky, appearing fast. Flowers (dark) yellow. Fruits 
orange-red, with sticky exudate; aril reddish. 


2. In Sinclair’s conception this species contains a number of specimens which are 
in the present revision regarded as a different species, H. affinis. The latter differs 
in many characters, such as the pedicel being articulate at base, a differently shaped 
androecium, a larger glabrous perianth, a different tomentum on the lower leaf 
surface, the perianth being persistent under the fruit, etc. 


3. H. motleyi, especially in a young stage, may be confused with Endocomia 
rufirachis, formerly Horsfieldia macrocoma var. rufirachis, a species also with 
pubescent flowers. 


61. Horsfieldia tomentosa Warb. Fig. 1C(61) 


Horsfieldia tomentosa Warb., Mon. Myrist. (1897) 302; Sinclair, Gard. Bull. Sing. 16 (1958) 403, fig. 40; 
28 (1975) 149 — Myristica tomentosa Hook. f. & Th., Fl. Ind. (1855) 161, nom. illeg., not of Thunberg 
(1782); A. DC., Prod. 14, 1 (1856) 204; Miq., Fl. Ind. Bat. 1 (2), 1 (1858) 68; Hook. f., Fl. Br. Ind. 5 
(1886) 105; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 301, pl. 129 — Type: Wallich Cat. 9025 
(‘“‘Myristicea?’’) (K-Wall.; iso: K, BM; CAL, G, SING, n.v.). 


Tree 5-20(-40) m. Twigs terete, towards apex 2-5(-10)) mm diam., late- 
grabescent from dense, rusty tomentum of woolly hairs c. 1.0-1.5 mm, bark of older 
twigs striate, not flaking; lenticels usually abundant, rather conspicuous. Leaves in 
2 rows, membranous, elliptic or obovate to oblong-lanceolate, broadest at + or 
slightly above the middle, 9-27 x 4-10(-12) cm, base nearly rounded to attenuate, 
tip acute-acuminate; upper surface drying brown, faintly minutely pustulate or not, 
glabrous or glabrescent, lower surface with persistent tomentum of dense dendroid 
hairs all of about the same size, c. 0.1-0.8 mm long, without blackish larger dots; 
midrib above flattish, glabrescent; nerves 7-15 pairs, flattish or sunk above, the 
marginal arches on lower surface rather distinct, rather regular of shape; tertiary 
venation forming a lax network usually indistinct or invisible above; petiole 10-18 x 
1.5-3 mm, densely pubescent; leaf bud narrowly ovoid-ellipsoid, c. 1-1.5 cm long, 
densely pubescent with hairs c. 1.5 mm. Inflorescences densely pubescent with 
woolly hairs 1.5-2.0 mm long, in CG’: rather many-flowered, 3 or 4 times ramified, 
3-12 x 1.5-7 cm, common peduncle 3-23 mm; in 9: + few-flowered, 2-6 cm long; 
bracts elliptic, densely woolly pubescent, 2-4 mm long, caducous. Flowers in small 
fascicles, 3-(or 4- or 5-) valved, glabrous; pedicels slender, at base not articulated, 
in O glabrous, in Q late-glabrescent, hairs c. 0.2 mm. Male perianth globose, 
1.4-2.5 mm diam., base and apex rounded; pedicels 1-3 mm long; perianth at 
anthesis cleft to nearly ’2-way, valves c. 0.1 mm thick. Androecium much- 
depressed globose, above flattish or usually impressed in the centre, circular to 
blunt-triangular in transverse section, 0.6-0.9 x 1.2-1.7 mm; anthers 9-12(-157), 
almost completely sessile, incurved towards the top; column broad, solid except for 
apical hollow c. 0.2 mm deep; androphore rather narrow, c. 0.4-0.5 mm long. 
Female perianths broadly ellipsoid, c. 2.5 x 2.3 mm , cleft at anthesis to c. %, 


I 


New account of Horsfieldia 3 211 


valves c. 0.2 mm thick: pedicel 1-2 mm long, late-glabrescent; ovary subglobose. c. 
1.5 mm diam., appressed-pubescent by hairs 0.1-0.2 mm, stigma minute, faintly 
2-lobed, c. 0.1 mm long. Fruits 1-5 per infructescence, ellipsoid, top and base 
rounded, 1.5-2.0 x 1.3-1.6cm, glabrescent, usually with minute tomentum remain- 
ing towards the base. drying dark brown, without lenticels but usually minutely 
pustulate, dry valves c. 1.5 mm thick; stalk 2-6 mm; perianth not persisting. 


Distribution. S. Peninsular Thailand, Malaya (Kedah, Penang, Perak, Treng- 
ganu, Pahang, Selangor, Malacca, Johore. Kelantan), Singapore (doubtful. see 
Sinclair, l.c. p. 150), Sumatra (E. Coast, fide Sinclair; no specimens seen). 


THAILAND. S. Peninsular: Geesink 7229, A.F.G. Kerr 17252, 17482; Phusomsaeng 142, 424 (BKF 
51976); Smitinand 10378. 


MALAYA. Cantley 30; Cuming 1846; Curtis 1197, 1748; Derry 967; FRI 0880, 3408, 15957, 16300. 
25099; Gaudichaud s.n., 44; Hb. Hooker s.n.; KEP 94965, 98507, 99205, 99403, 104304; King’s Coll. 
4165, 5671, 6102, 8532, 8642, 10557; Shah (& Noor) MS. 1319, 1536, 1864; Phillips s.n.; Ridley s.n., 44. 
7205, 10240; SFN 1082, 21751, 35134; Sinclair 9878, 10158; Unesco Limestone Exp. 159; Wall. Cat. 9025. 


Ecology. Lowland and foot hill forest, in S. Thailand in evergreen forest, old 
secondary forest; 0-300 m alt. Flowers mostly in March, fruits mostly in July. 


NOTES 


1. Fieldnotes. Small or moderate, sometimes big tree to c. 40 m. alt. Bark 
recorded as fissured, not flaky or scaly, brown to blackish, soft. Inner bark pale 
reddish or pale yellowish, laminated, fibrous; exudate pink-red: sapwood whitish 
or pink. Flowers yellow, with a fine perfume. Fruits yellow to orange, aril red: 
perianth persistent in fruit. 


2. The lobes of the male perianths in fully mature flowers are often slightly 
outward recurved. 


3. H. tomentosa is in the present revision accepted in the same sense as by 
Sinclair and earlier authors. It belongs in the alliance with H. grandis, H. flocculo- 
sa, H. motleyi. 


4. The name tomentosa should be attributed to Warburg because Myristica 
tomentosa Hook. f. and Thoms. is illegitimate; being a later homonym. 


62. Horsfieldia gracilis de Wilde, sp. nov. 


Horsfieldia paucinervis Warburg differt habitu gracili, foliis tenuiter membranaceis, tomento persist- 
ente, nervis paribus 14-17, fructibus ellipsoideis, c. 1.5 cm longis, glabris, perianthio persistente. — 
Type: Sarawak, Ilias Pai’e S 16604 (L: iso: K; S, n.v.). 


Tree c. 5 m. Twigs terete, towards the apex 1.5-2.5 mm diam., rather late 
glabrescent, tomentum light brown with stellate-dendroid hairs 0.3-0.5 mm, bark of 
older twigs striate, neither cracking nor flaking; lenticels minute and inconspicuous 
or absent. Leaves in 2 rows, thinly membranous, oblong-lanceolate, broadest at 
about or slightly above the middle, 12-21 x 4-6(-6.5) cm, base (rounded-) 
attenuate, tip acute-acuminate; upper surface drying dull olivaceous, glabrous, 
lower surface pale olivaceous-brownish, with subpersistent tomentum consisting of 
scattered pale dendroid-stellate hairs 0.3(-0.5) mm; without larger brownish or 
blackish dots or marks: midrib raised above, beneath with persistent tomentum; 


212 Gard. Bull. Sing. 38(2) (1985) 


nerves 14-17 pairs, slightly raised to flat above, the marginal arches regularly 
shaped, slightly impressed and obvious; tertiary venation forming a lax network, 
indistinct; petiole 8-14 x 1.5(-2) mm, late glabrescent; leaf bud c. 6-7 X 1.5 mm, 
densely pale-brown pubescent, with hairs 0.3-0.5 mm long. Male flowers not seen. 
Female inflorescences not or only once ramified, 2-5-flowered, 1-1.5 cm long, 
pubescent with hairs 0.3-0.5 mm long; bracts not seen. Female flowers 3-valved, 


glabrous, pedicel at base apparently inarticulate (male flowers not seen); perianth — 


(persistent as judged from the fruit) c. 2.0 x 1.5 mm, at anthesis cleft to c. 2-way, 
valves c. 0.3 mm thick, glabrous; pedicel c. 1 mm long, glabrous; pistil not seen. 
Fruits 1-3 per infructescence, ellipsoid, top and base subobtuse, 1.4-1.5 x 1.0-1.1 
cm, glabrous, drying dark brown, with a finely granulate structure, lenticel-like 


tubercles absent, dry valves 0.5-1.0 mm thick; stalk 1-1.5 mm; perianth persistent. 


under the fruit. 
Distribution. Borneo: Sarawak, Miri Dist., only known from the type. 
Ecology. Primary lowland forest. 
NOTES 
1. This species is known only from the type collection, in fruit. It was recorded as 
a small tree, 15 feet high and 4 inches in girth, and identified by Sinclair as 
Horsfieldia sp. Although only the fruits are known, it is obvious from its general 


habit that the present new species belongs to the group of H. paucinervis, beside 
which it keys out. It is inferred that the male flowers have the following characteris- 


tics: inflorescences rather tiny, c. 6 cm long, pubescent with hairs c. 0.4 mm long;. 


perianth subglobose (likely c. 1-1.5 mm diam.), 3-(or 4-)valved, splitting in bud to 
c. 2-way, glabrous; pedicel glabrous, at base inarticulate; androecium globose or 
depressed-globose, sessile or with a short androphore, in transverse section + 
circular, anthers largely sessile, the column at the apex with a shallow hollow. 


Horsfieldia gracilis is readily recognized by its slender habit, thin membranous 
leaves and (sub)persistent though not very conspicuous, rough tomentum on the 
leaf bud, twig apex and lower leaf surface, especially on the midrib and nerves, and 
by the small fruits with a persistent perianth. Superficially the species may be taken 
for H. tenuifolia or H. macilenta. 


63. Horsfieldia paucinervis Warb. Fig. 1C(63) 


Horsfieldia paucinervis Warb., Mon. Myrist. (1897) 345, t. 22; Merr., En. Born., J. Str. Br. R. As. Soc., 
spec. number (1921) 268; Sinclair, Gard. Bull. Sing. 28 (1975) 93 — Myristica paucinervis (Warb.) 
Boerl., Handl. Fl. Ned. Ind. 3.1(1900) 87 — Type: Sarawak, Beccari 3279 (Q fl.) (FI, n.v.). 


Tree 3-8 m. Twigs terete, 2-3 mm diam., late glabresent, tomentum reddish to 
yellow-rusty, of shaggy hairs 1.5-2 mm long, bark of older twigs dark grey, striate, 
not cracking; lenticels not conspicuous. Leaves in 2 rows, thinly chartaceous, not 
bullate, elliptic to elliptic-oblong, broadest at about the middle, 7-15 x 3-6.5 cm, 
base rounded to attenuate, tip acute to acute-acuminate; upper surface drying 
olivaceous to brown, glabrous, lower surface dull brown with persistent, thin 
tomentum of dendroid hairs of mixed sizes, c. 0.5-0.1 mm long; larger blackish dots 
or stripes present (always?); midrib flat above, indistinct, with persistent tomentum 
or late glabrescent; nerves 5-9 pairs, flat or sunken, the marginal arches indistinct; 


, oe — 


-e 


New account of Horsfieldia 3 213 


tertiary venation very indistinct or invisible; petioles 6-12 X 1.5-2.5 mm, pubescent; 
leaf bud c. 8-10 x 3-4 mm, long pubescent. Inflorescences densely woolly pubes- 
cent with hairs (0.7-) 1.0-2.0 mm long, in CG’: many-flowered, 3 or 4 times ramified, 
5-9 x 3-5 cm, common peduncle up to 15 mm long; in 9: 4-6 cm long; bracts not 
seen, caducous. Flowers in © in loose clusters, 3-valved, perianth glabrous, 
pedicels glabrous, at base inarticulate. Male perianth subglobose, 0.8-1.0 x 1.0-1.4 
mm, top and base broadly rounded; pedicels slender, 0.5-1.0 mm long; perianth at 
anthesis cleft to c. 4 to nearly '12-way, valves c. 0.1 mm thick. Androecium small, 
+ depressed-globose, c. 0.3-0.5 x 0.4-0.8 mm, + circular in transverse section (not 
3-angled); anthers 4 or 5 (thecae 8 or 10), almost completely sessile, the tips 
incurved; column broad, solid except for a minute hollow at the top c. % deep, 
androphore narrow, somewhat tapering, c. 0.1-0.2 mm long. Female perianth 
(according to Warburg, Sinclair) ovoid-globose, c. 2.0 mm diam., cleft at anthesis 
to somewhat over '2-way; pedicel c. 2 mm long, glabrous; ovary subglobose, 
glabrous, stigma minute. Fruits (immature) with persistent perianth, oblong, glab- 
rous, obtuse at both ends, c. 1.0 x 0.7 cm; stalk c. 2 mm long. 


Distribution. Borneo: Sarawak (1st, 2nd, 4th Divisions). 


BORNEO. Sarawak: Beccari 3279, n.v.; Haviland 1735, 3075; Purseglove P 4403. 


Ecology. Coastal kerangas and secondary forest on eroded white sand; 0-50 m. 
Flowers throughout the year. 


NOTES 


1. Fieldnotes. Shrub or slender tree to c. 8 m tall. Sap pale pink, watery. Flowers 
yellow. 


2. Apparently a member of the group of which leaves are pubescent and to 
which also belong H. splendida, H. reticulata, H. rufo-lanata and H. tomentosa, and 
somewhat more remotely, H. grandis. H. paucinervis is readily recognized by its 
few nerves, almost invisible reticulation and small male flowers, with few anthers. 


3. I have seen only a few male flowering specimens, incl. Haviland 3075 and 
Purseglove P 4403 but not the type which is Beccari 3279, a female flowering 
one. This last is described and depicted by Warburg; Sinclair apparently also saw a 
specimen with immature fruits. It is odd that ovaries and fruits are reported as 
glabrous as they are pubescent in most related species though glabrous in H. 
motleyi. 


64. Horsfieldia splendida de Wilde, sp. nov. Fig. 1C(64); 24 


Ramuli validi, tarde glabrescentes, pilis 1-1.5 mm longis obtecti, foliis + oblongis, 18-45 cm longis, 
subtus tomento persistente. Perianthium masculum subglobosum, 1.5-2 mm diam., 3- vel 4- valvatum, 
androecio depresso-globoso, apice moderate excavato, antheris 8-10, sessilibus, androphoro angusto, 
0.3-0.5 mm longo, pedicello gracili, basi non-articulato, fructibus late ellipsoideis, c. 2 cm longis, 
pubescenti, perianthio persistente. — Type: Sarawak, Othman Ismawi S 33723 (L; iso: NO, SAN, 
n.v.. 


Tree 10-20(-30) m. Twigs stout, terete, towards apex 4-7(-13) mm diam., late 
glabrescent, tomentum densely woolly, yellow-brown to rusty, with hairs 1.0-1.5 
mm, bark of older twigs usually dark brown or blackish, coarsely longitudinally 
striate and fissured, later on cracking and + flaking; lenticels few, indistinct. 


Sais ae: 
i : 


Y 
Ys 
fi 
SE SAAS 


\ 


FARSI SSF ENS. tteme an StS, 
q \ 


Fig. 24. Horsfieldia splendida de Wilde. 
a. apical part of leafy twig, x 2; b. portion of twig with immature male inflorescence axillary 
to leaf scar, note bracts, x 42: c. mature male inflorescence, x 2; d. mature male flower, 
lateral view, x 12; e. ditto, longitudinally opened, showing androecium, x 12; f. androecium, 
longitudinal section, schematic, x 12; g. portion of twig with infructescence, fruits immature, 
x 2; h. mature fruit, note persistent perianth, x /2.—a, b from Anderson 12916; c-f. from 
B.N.B. For. Dept. 4782; g. from San. 16927; h. from SFN 35606. 


214 


New account of Horsfieldia 3 25 


Leaves in 2 rows, membranous to chartaceous, elliptic-oblong to oblong- 
oblanceolate, broadest usually above the middle, 18-45 x 6.5-17 cm, base almost 
rounded to attenuate, tip acute-acuminate; upper surface drying dull olivaceous to 
brown, not minutely pustulate, glabrous (early glabrescent), lower surface with 
conspicuous rather dense tomentum of loosely branched, stalked dendroid hairs, + 
evenly spaced and of about the same size, hairs 0.5-1.5 mm long; without larger 
blackish dots; midrib above late glabrescent, flat or slightly raised; nerves 18-25 
pairs, above sunken, early glabrescent, the marginal arches usually distinct and 
regularly shaped; tertiary venation forming a lax network, generally distinct above; 
petioles 12-17 x 3.5-5 mm, pubescent; leaf bud c. 15(-20) x 5 mm, with hairs 1-1.5 
mm long. Inflorescences densely woolly pubescent with dendroid hairs 1-1.5 mm, 
in ©: many-flowered, 4(or 5) tumes ramified, 6-16 x 3-12(-16) cm, common 
peduncle 5-20 mm; Q inflorescences rather few-flowered, 3-5 cm long; bracts + 
ovate-triangular, acuminate, 0).4-1 cm long, densely pubescent, caducous. Flowers 
in © in loose clusters, 3-(or 4)-valved, glabrous or thinly pubescent, pedicels 
slender, glabrous or subglabrescent, hairs 0.3-0.5 mm long, at base inarticulate. 
Male perianths subglobose, usually somewhat depressed, 1.4-2 x 1.6-2 mm, top 
broadly rounded, pedicels 1-1.5 mm long, slender; perianth at anthesis cleft to to 
nearly '/2-way, valves c. 0.2 mm thick. Androecium depressed-subglobose, usually 
+ impressed in the centre, circular or faintly 3-angular as seen from above, 0.5-0.6 
x 1-1.3 mm; anthers 8-10, completely sessile, incurved towards the top; column 
broad, solid, leaving a small apical hollow of c. ¥%3-(2-way) deep; androphore 
rather slender, 0.3(-0.5) mm long. Female perianth not seen, according to the 
persistent remnants under the fruit c. 3.0 x 2.5 mm, 3- or 4-valved, outside 
pubescent; pedicel 0.5-1.0 mm long, pubescent; ovary ovoid-ellipsoid, pubescent; 
stigma minutely 2-lobed, c. 0.1 mm high. Fruits 2-7 per infructescence, broadly 
ellipsoid, obtuse at apex, base rounded, 2.0-2.2 x 1.5-1.8 cm, pubescent (hairs c. 
0.5 mm), drying brown, without lenticels or tubercles, dry valves c. 3 mm thick; 
stalk c. 1 mm long; perianth persistent. 


Distribution. Borneo. Sarawak, Brunei, Sabah, E. Kalimantan; probably W. 
Kalimantan (see notes). 


BORNEO. Sarawak: Anderson 12916, 15526; Galau (tree nr Semengo For. Res) 3500; Motley in Hb. 
Hooker 178; Zen Osman 5140; S§ 33723; Seal 547; SFN 35606; Sinclair & Kadim 10227 —Sabah: B.N.B. 
For. Dept. 4782; SAN 15390, 15487, 16656, 16927, 23739, 36333, 36592, 36660, 44688, 73393, 74992, 
78355; Sinclair (Kapis & Kadim) 9292 — E. Kalimantan (W. Kutai): Endert 5010. 


Ecology. Lowland mixed Dipterocarp forest, edges of swamp forest, kerangas 
forest (with Dryobalanops fusca dominant), also montane forest; brown soil, 
yellowish soil, tuff-plateau; 0-600(-1500) m alt. Flowers and fruits throughout the 
year. 


NOTES 


1. Fieldnotes. Bark of trunk usually recorded as blackish, brown, or red-brown 
and fissured or flaking. Inner bark soft, light brown or reddish, laminated, exudate 
reddish. Wood yellowish or whitish, recorded as soft or medium hard. Flowers 
yellow, fragrant (like the odour of Peru-balsam). Fruits recorded as red or orange. 


2. The present new species contains the majority of the specimens which were 
determined as H. reticulata by Sinclair. However, the type of H. reticulata, Beccari 
3475 (FI, n.v.; iso: K) clearly represents a different species, one with the male 


216 Gard. Bull. Sing. 38(2) (1985) 


pedicels articulate at base, much smaller, more coriaceous, more distinctly reticu- 
late leaves, and a much less pronounced indumentum on the lower surface. Its 
flowers, however, are almost similar, only a trifle larger. See further under that 
species. 


3. I have not seen Hans Winkler 388 (HBG), from West Kalimantan, deter- 
mined by Sinclair as H. reticulata; may be it is our present new species H. 
splendida. 


4. H. splendida is closely related to H. reticulata, H. rufolanata, H. tomentosa, 
and also H. grandis and less so to H. flocculosa; all these having similarly shaped 
and constructed male flowers. The perianth remains persistent under the fruit in 
these species except in H. tomentosa and H. rufolanata. 


65. Horsfieldia rufo-lanata Airy-Shaw Fig. 1C(65) 


Horsfieldia rufo-lanata Airy-Shaw, Kew Bull. 10, 1939 (1940) 440; Sinclair, Gard. Bull. Sing. 28 (1975) 
111 — Type: Richards 1667 (K). 


Tree 8-17 m. Twigs terete, towards the apex 3.5-5(-7) mm diam., late glabres- 
cent, tomentum dark rusty to reddish-brown, of hairs 1-1.5 mm long, coarsely 
striate, bark of older twigs dark brown, + longitudinally cracking, not flaking; 
lenticels present but indistinct. Leaves in 2 rows, chartaceous, slightly or not 
bullate, elliptic to oblong, broadest at the middle, 10-23 x 5-10(-12) cm, base 
nearly rounded to subattenuate, tip acute to acute-acuminate; upper surface drying 
olivaceous to brown, glabrous except the nerves, lower surface with dense to sparse 
tomentum of rather harsh hairs of mixed sizes ranging from sessile-stellate, c. 0.3 
mm, to short-armed, dendroid emerging hairs up to c. 1.5 mm long; without larger 
blackish dots; midrib above + raised, late glabrescent; nerves 11-16 pairs, raised, 
pubescent or late glabrescent, the marginal arches distinct and regularly shaped; 
tertiary venation forming a lax network, sunken, distinct or not; petioles 10-16 x 
3-4.5 mm, pubescent; leaf bud 15-20 x 3-5 mm with hairs 1-1.5 mm. Inflorescences 
densely woolly-shaggy pubescent, hairs 2.0-3.0 mm long, in C’: many-flowered, 3 
or 4 times ramified, 8-12 x 5-7 cm, common peduncle 15-30 mm; in Q: few- 
flowered, c. 3 cm long; bracts densely shaggy pubescent, oblong-triangular to 
lanceolate, 3-8 mm long. Flowers in OC in loose clusters, 3- or 4-valved, perianth 
glabrous, pedicel glabrous, at base indistinctly articulate. Male perianths subglo- 
bose, hardly or not depressed, c. 2 x 2.3 mm, rather firm; pedicels not very 
slender, c. 0.5 mm thick, 0.5-1 mm long; perianth at anthesis cleft to nearly ¥2-way, 
valves 0.2(-0.3) mm thick. Androecium globose (not depressed-globose), c 1.2 mm 
diam., not or only slightly apically impressed, circular in transverse section; anthers 
c. 15 (c. 28-30 thecae, very closely appressed), completely sessile, incurved towards 
apex; column broad, solid except for the small apical hollow to c. %, deep; 
androphore slender, c. 0.2 mm long. Female perianth not known. Fruits 2-4 per 
infructescence, broadly ovoid-ellipsoid, top and base rounded, 2.0-2.2 x 1.5-1.6 
cm, glabrescent, remnants of minute hairs c. 0.1 mm at the base, drying brown, 
without distinct lenticels or tubercles, dry valves c. 2 mm thick; stalk 1-3 mm long; 
perianth not persisting in mature fruit. 


Distribution. Borneo: Sarawak (4th & Sth Div.), Sabah. 


BORNEO. Sarawak: (Briinig) S 10599, n.v.; Richards 1667 — Sabah: (Wood) SAN 16295; (Meijer) 
26423. 


New account of Horsfieldia 3 217 


Ecology. Montane forest, upper Dipterocarp forest on sandstone; 900-1400 m 
alt. Flowers in June and September, fruits in September. 


NoTES. A montane species closely related to H. splendida and especially H. 
reticulata. Differs by the longer woolly tomentum of the inflorescences, the almost 
globose, hence not depressed-globose, male perianth, the near-globose 
androecium, more anthers, 13-15(-20) as against 8-10 and 10-12 in H. splendida and 
H. reticulata respectively. The nerves on the upper leaf surface apparently remain 
pubescent for a much longer time, and are generally raised. In contrast with H. 
splendida, the blades are generally smaller and more elliptic and have a tomentum 
of stiffer hairs on the lower surface; H. reticulata has a much less conspicuous 
tomentum on the lower leaf surface. 


66. Horsfieldia affinis de Wilde, sp. nov. Fig. 1C(66) 


Folia membranacea, in sicco fusco-brunnea, subtus pilis persistentibus 0.5-1 mm longis obtecta. Flores 
masculi glabri, perianthio depresso-globoso, 1.2-1.5 Xx 1.5-2.0 mm, 3 vel 4 -valvato, androecio 
subgloboso, c. 1 mm diam., apice excavato usque ad 2, antheris 11, sessilibus, pedicello tenere, basi 
articulato, fructibus ellipsoideis, c. 2.5 cm longis, glabris, perianthio persistente. — Type: Sarawak, 
E. Wright S 24718 (L; iso: K). 


Tree 7-15 m. Twigs terete, towards the apex 3-6(-9) mm diam., late-glabrescent, 
tomentum of rusty hairs 0.5-1.0 mm long, lower down with the bark finely striate, 
when older not flaking; lenticels absent or small and inconspicuous. Leaves in 2 
rows, membranous, not bullate, elliptic-oblong to lanceolate, broadest at or below 
the middle, 18-35 x 5-11 cm, base rounded to attenuate, top acute-acuminate; 
upper surface drying bright to dark or blackish brown, glabrous, lower surface with 
persistent, rather sparse tomentum of dendroid hairs or mixed with sessile stellate, 
dendroid hairs 0.5-1.0 mm long; scattered larger blackish dots absent; midrib 
flattish or slightly raised above, often late glabrescent; nerves 17-20 pairs, slender, 
raised, glabrous above, the lateral arches usually regularly shaped and distinct; 
tertiary venation forming a lax network, rather faint to distinct above; petioles 
12-20 x 2-4 mm, pubescent; leaf bud rather slender, 10-15 x 3-4 mm, densely 
pubescent with hairs 0.5-1 mm. Inflorescences densely pubescent with rather 
shaggy hairs 0.5-2.0 mm long, in CO’: many-flowered, c. 4 or 5 times ramified, 12-25 
x 8-14 cm, common peduncle 15-60 mm long; @ inflorescences 5-6 cm long, 
fewer-flowered than the males; bracts broad-triangular to elliptic-oblong, acute, 
3-9 mm long, densely pubescent, caducous. Flowers in C in loose clusters of 2-6 
each, 3- or 4-valved; perianth glabrous, pedicel slender, glabrous, at base inarticu- 
late. Male perianth depressed-globose, 1.2-1.5 x 1.5-2.0(-2.2) mm, top and base 
broadly rounded; pedicel slender, 1.5-2 mm long; perianth at anthesis cleft to c. 
Y4-¥3; valves c. 0.1 mm thick. Androecium subglobose or depressed globose, 
slightly impressed at the centre, + circular in transverse section, c. 0.7-0.8 x 
1.0-1.2 mm; anthers 11, completely sessile, incurved towards the apex; column 
broad, solid except the apical hollow up to c. ’2-way deep; androphore slender or 
rather broad, 0.2-0.3 mm long, hidden or not by the anthers. Female flowers not 
seen; perianth as judged from remnants under the fruit 2.5-3.0 < 2-2.5 mm, valves 
3, splitting the bud to over 2-way, glabrous or probably with very weak tomentum. 
Fruits 4-7 per infructescence, ellipsoid, top and base rounded, 2.3-2.7 x 1.7 X 2.1 
cm, glabrous, drying brown, without lenticels but paler, wart-like tubercles pre- 
sent, dry valves 1.5-2.0 mm thick; stalk 3-4 mm long; perianth persistent under the 
fruit. 


Distribution. Borneo: Sarawak; Brunei; Central, East and SE. Kalimantan. 


218 Gard. Bull. Sing. 38(2) (1985) 


BORNEO. Sarawak (Kapit, 3rd Div.): Wright S 24718; (Chai) S 36018 — Brunei: (Abdul Latip) 
BRUN 5654 — Kalimantan, East & SE.: Endert 2595; Kostermans 9535, 21760; Leighton 700, 737 930; 
Sauveur I 14, 126; Central: Nooteboom 4847. 


Ecology. Forest on alluvial soils, riverbanks: sandy clay soil, sandy ridges, 
sandstone; 0-600 m alt. Flowers May to November, fruits in July. 


Vernacular name. Kumpang balau (Iban name, Kapit area, Sarawak). 
NOTES 


1. Fieldnotes. Small tree, crown pyramidal, no buttresses. Bark rough, dark 
brown, lenticellate; outer bark 1-2 mm thick, brown-red, inside red. Flowers dark 
or golden yellow, fragrant or with strong sweet smell of Peru-balsam. Fruits 
ramiflorous, in bunches behind the leaves, pinkish-orange. 


2. Most specimens of the present new species were included by Sinclair in H. 
motleyi, which differs by the inarticulate pedicels, the much smaller pubescent 
flowers with a quite different androecium, the leaves drying to a dull olivaceous 
brown colour, and the sunken lateral nerves. 


3. H. affinis is among its closest relatives characterized by the distinctly articu- 
late, glabrous, male pedicels, the male flowers being arranged in small clusters at 
the end of the slightly thickened hairy inflorescence-ramifications. A second re- 
lated species with articulate pedicels is H. reticulata, and possibly H. rufo-lanata the 
position of which is unconfirmable because there are not enough flowers available. 


67. Horsfieldia reticulata Warb. Fig. 1C(67) 


Horsfieldia reticulata Warb., Mon. Myrist. (1897) 304, t. 22 fig. 1-3; Merr., En. Born. J. Str. Br. R. As. 
Soc. special number (1921) 268; Sinclair, Gard. Bull. Sing. 28 (1975) 107 — Myristica reticulata 
(Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 85 — Type: Sarawak, Beccari 3475 (FI, n.v.; iso: K). 


Tree to c. 20 m. Twigs terete, towards the apex 2.5-4(-8) mm diam., late 
glabrescent, tomentum rusty, of rather coarse hairs c. 1.0 mm long, bark of older 
twigs dark grey, coarsely striate, later on finely longitudinally and transversely 
cracked; no lenticels seen. Leaves in 2 rows, chartaceous, + bullate, elliptic to 
oblong-oblanceolate, broadest at or above the middle, 8-24 x 4-6.5 cm, base 
rounded, top acute-acuminate; upper surface drying blackish brown, glabrous, 
lower surface bright dark brown, with persistent tomentum composed of rather 
spaced hairs of various sizes, 1.e., stellate sessile hairs, and long-emergent short- 
armed dendroid hairs up to c. 0.7 mm long as well as intermediate forms; larger 
blackish dots absent; midrib raised above, glabrous; nerves 9-19 pairs, sunken, 
glabrous, the marginal arches distinct and regular; tertiary venation forming a lax 
network, very distinct on both surfaces; petioles 7-15 x 2-3 mm, pubescent; leaf 
bud not seen. Inflorescences in CG’: on the older wood, densely pubescent with hairs 
0.5-1.0 mm, many-flowered, c. 4 times ramified, c. 18 x 12 cm, common peduncle 
20-25 mm; bracts not seen, caducous. Flowers 3(-5)-valved in small loose clusters; 
perianth glabrous; pedicels glabrous, at base articulate. Male perianth somewhat 
depressed-globose, 1.5-2.0 x 2.0-2.5 mm, top and base broadly rounded; pedicel 
slender, 0.5-1.5 mm long; perianth at anthesis cleft to c. “4 to nearly 2-way; valves 
0.1-0.2 mm thick. Androecium much depressed-globose, impressed in the centre, 
c. 0.6-0.7 x 1.2-1.5 mm, in transverse section + ellipsoid or faintly 3-angular; 
anthers c. 10-12 or c. 15 (see notes), + completely sessile, incurved towards the 


~~ Dee 


New account of Horsfieldia 3 219 


top; column broad, solid except for an apical hollow with flattish bottom reaching 
to depth of c. % to nearly '’2-way; androphore rather slender, 0.3-0.4 mm long, 
largely hidden by the anthers. Female flowers and fruits not seen. 


Distribution. Borneo: Sarawak (2nd. Div.), East & West Kalimantan. 


BORNEO. Sarawak (2nd Div.): Beccari 3475 — Kalimantan: West, Hans Winkler 388; East, 
Leighton 655 (possibly, material incomplete). 


Ecology. Forest at c. 100 m. alt. Flowers in November. 


NOTES 
1. Fieldnotes. Flowers yellow. Fruits sourish, eaten by the Dajaks. 


2. This species appears to be represented only by the type and two collections 
from West-Central and East Kalimantan. Most specimens considered by Sinclair as 
conspecific (Sinclair, p. 107, 108) belong to a presently described new species, H. 
splendida. Sinclair observed that the Beccari specimen looked somewhat different, 
but regarded it as a relatively glabrous specimen from which most tomentum had 
dropped off. H. splendida differs, in addition to the inarticulate pedicels, by the 
much less bullate and larger leaves, by the tomentum (of the lower leaf surface) of a 
different structure; the dendroid hairs being more widely branched, all + evenly 
long-stalked, and by its somewhat smaller male perianth. It should be noted, 
however, that the male flower including the androecium, is much the same in all 
species of the group of obviously related species to which H. grandis, H. tomentosa, 
H. flocculosa also belong. Because of the articulate pedicels, H. reticulata seems 
most related to H. affinis. 


3. The general habit of the leaves and inflorescences of the few collections seen 
by me is very much the same, the leaves being brittle-chartaceous, drying dark 
brown, with a tomentum rather harsh and a very pronounced reticulation which is 
impressed above. The last gives the leaves a distinct bullate appearance, more so 
because the leaf-margin is revolute. The androecia of the two male flowering 
specimens seen, however, are rather different. In Beccari 3475 I counted (9 or) 
10-12 anthers, in Winkler 388 there are apparently c. 15 anthers (c. 30 thecae), and 
the apical hollow of the column in the collection is generally broader, and has a 
wide, flattish bottom. 


68. Horsfieldia crassifolia (Hook. f. & Th.) Warb. Fig. 1C(68); 25 


Myristica crassifolia Hook. f. & Th., Fl. Ind. (1855) 160; A. DC., Prod. 14, 1 (1856) 204; Miq., Fl. Ind. 
Bat. 1 (2), 1 (1858) 68; Hook. f., Fl. Brit. Ind. 5 (1886) 108; King., Ann. Roy. Bot. Gard. Calc. 3 
(1891) 308, pl. 140 — Horsfieldia crassifolia (Hook. f. & Th.) Warb., Mon. Myrist. (1897) 323 (p.p); 
Sinclair, Gard. Bull. Sing. 16 (1958) 386, fig. 34, Pl. X A: 28 (1975) 23; Anderson, Gard. Bull. Sing. 
20 (1963) 195 — M. irya var. crassifolia Miq. ex Hook. f., Fl. Brit. Ind. 5 (1886) 108, pro syn. — Type 
Griffith 4350 (K; iso: CAL, n.v.) (see notes by Sinclair, l.c. pag. 25). 


M. horsfieldia auct. non. Bl.: Wall. Cat. (1832) No 6806, p.p. (other parts are H. polyspherula and H. 
wallichii). 


M. subglobosa Migq., Fl. Ind. Bat. Suppl. 1, 3 (1861) p.p. (other part is H. irya). 
M. paludicola King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 328, pl. 169 — H. fulva (King) Warb. var. 


paludicola (King) Warb., Mon. Myrist. (1897) 299 — Type: King’s Coll. 4267 (CAL, n.v.; iso: L, 
BM), 4706 (CAL, n.v.; iso: L, K, BM), 6688 (CAL, n.v.; iso: K); Wray 3071 (CAL, n.v.; iso: K, L). 


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Horsfieldia crassifolia (Hook.f. & Th.) Warb. 

a. branch with leafy twig and infructescence with mature fruits, note persistent perianth and 
aril completely closed, x 2; b. portion of twig with male inflorescence, x 12; c. mature male 
flower, lateral view, x 12; d? ditto, longitudinally opened, showing androecium, X 12; e. 
portion of older twig with female inflorescence axillary to leaf scar, x 2; f. mature female 
flower, opened, showing glabrous ovary and 2-lipped stigma, note flower considerably larger 
than male flower, X 12; g. portion of lower leaf surface with persistent tomentum and 
irregularly shaped dark-coloured dots, X 25. —a. from Mondi 51; b-d. from S 9226; e-g. from 
San. 27183. 


New account of Horsfieldia 3 221 


Tree 10-25 m. Twigs terete, not ridged, towards apex 2-6(-8) mm diam., rather 
early glabrescent, tomentum yellow-brown to rusty woolly, with hairs 0.2-0.5 mm; 
bark coarsely striate, when older not flaking, lenticels sparse to dense, distinct or 
not. Leaves in 2 rows, coriaceous, elliptic to oblong, 10-20(-28) x 3.5-7(-10) cm, 
base rounded to attenuate, top rounded to subacute or rarely emarginate; upper 
surface drying dull greenish brown to dark brown, not finely pustulate, lower 
surface either largely or partly covered with subpersistent tomentum consisting of 
interwoven stellate scales c. 0.1 mm high (when very young with dendroid emer- 
gents), or glabrescent and then showing distinct hair scars and usually scattered 
blackish dots or dashes; midrib above flattish; nerves 11-16 pairs, above thin and 
flattish or sunken; tertiary veins indistinct or invisible on both surfaces; petiole 
9-20(-30) x 1.5-4.5 mm, leaf bud 7-12 2-3 mm with hairs 0.2-0.5 mm. Infloresc- 
ences situated in between or usually behind the leaves, late glabrescent or with a 
persistent, rather dense, woolly tomentum of dendroid hairs 0.2-0.5 mm; in CG: 3-5 
times ramified, broadly paniculate, many-flowered, 6-20 x 4-15 cm, common 
peduncle 5-20 mm; in Q: 3-14 cm long; bracts elliptic-lanceolate, 2-5(-7) mm long, 
pubescent, caducous. Flowers in loose clusters of 2-7, glabrous; perianths 2-valved; 
pedicels slender, glabrous, at base inarticulate. Male perianth globose or slightly 
transversely ellipsoid, (0.8-)1.0-1.3 x 1-1.5 mm, the upper part broadly rounded, 
the base broadly rounded to flattish, pedicel c. 0.3-1 mm long; perianth at anthesis 
cleft to c. Y3-2(-*%), valves 0.2-0.3 mm thick. Androecium globose or + transverse- 
ly ellipsoid, not or only faintly laterally compressed, c. 0.4-0.5 x 0.5-0.8 mm; 
anthers 4-6(thecae 8-12), + septate before maturity, widely spaced, and with broad 
connectives (giving the androecium an angular appearance), the anthers free for 
almost the upper half or more; androphore c. 0.2(-0.3) mm long, slender. Female 
perianth broadly obovoid, 2-3 xX 2-2.5 mm, split at anthesis for “5-3 only, valves 
0.6-1 mm thick; pedicel 1.5-2.5 mm long; ovary obovoid, glabrous, 1.5 x 1.2-1.5 
mm, stigma minute, consisting of 2 small lobes c. 0.1-0.2 mm high, running out into 
a faint ridge at one side of the ovary. Fruits (1-)2-10 per infructescence, ovoid to 
obovoid, base and top rounded, 1.5-2.2 x 1.2-1.8 cm, glabrous, drying dark brown, 
with few or no lenticel-like tubercles; dry valves 1.5-2 mm thick; stalk 2-5(-7) mm 
long; perianth persisting under the fruit. 


Distribution. Malaya (Perak, Trengganu, Selangor, Negri Sembilan, Malacca, 
Johore), Singapore, Sumatra (incl. Indragiri, Riau, Bangka & Belitung), Borneo; 
absent in S. Thailand. 


MALAYA. Anderson 9; Derry 1163; FRI 0021 7727, 17634; Hb. Griffith 4350; KEP 70486; King’s 
Coll. 4267, 4706, 6688; K.C. Liew 166; SFN 32105, 40567, 40898; Wray 3071. 


SINGAPORE. Ridley 1819, 1828, 3831, 5826, 6126, 6909, 10695; SFN 34542, 34905, 37710, 39533, 
40256. 


SUMATRA. Tapanuli: b.b. 28/69, 28399, 29545; Rahmat si Toroes 4829; Theunissen 59, 60 — W. 
Coast: b.b. 5952; Korthals s.n. — E. Coast: b.b. 21290 — Riau: b.b. 20375 — Bangka: Grashoff 117. 


BORNEO. Sarawak: Anderson 9028; Haviland & Hose 1941; § 9226, 12325, 32021; Sanusi bin Tahir 
5204, 9279; SFN 36078 — Brunei: Brun. 375, 838; Sinclair & Kadim 10473 — Sabah: B.N.B. For. Dept. 
10604; Kokawa & Hotta 326; SAN A 4562, 4598, 24313, 27131, 27183, 27826, 78031, 84511 — 
Kalimantan: West, Mondi 5] — East & SE.: b.b. 9937, 32404, Kostermans 4189, 9857A, 10097; de 
Vriese s.n. 


Ecology. Mostly in marshy forest, freshwater and peat swamp-forest; on sandy 
soils, c. 0-200 m alt. Flowers and fruits throughout the year. 


Vernacular names. Jangkang paya (Malay); Kajoe haroeja (Batak), Kumpang 
ensuliue, Kumpang sadara, Terada’a, Ta’dara (Sarawak, fide Anderson). 


$9 3, Gard. Bull. Sing. 38(2) (1985) 


NOTES 


1. Fieldnotes. A few stilt roots or low buttresses occasionally recorded. Flowers 
often recorded as yellow, strongly scented. Bark greyish, fissured, flaking in small 
rectangular scales. 


2. The lower surface of the leaves of Bornean material tend to become some- 
what earlier glabrescent as compared wtih those in Sumatra and Malaya. 


3. May be confused with H. fulva, a species also with + coriaceous leaves and a 
persistent perianth on the fruit, but this has a 3-merous perianth. Sterile specimens 
of the present species may be recognized by the coriaceous leaves, and on the lower 
surface by the usually pesistent ‘scaly’ tomentum, and the usually present sparse to 
rather dense irregularly-shaped dark-brown dots and streaks. Sinclair (1975, p. 26) 
remarks that the species can easily be recognized from a distance by its rusty or 
cinnamon-brown colour of the lower leaf surface. The species is very constant in 
habit, characters, and habitat. 


It has few (only c. 4-6) well separated anthers, the androecium being typically 
narrowly attached to the base of the perianth. 


It is one of the few West Malesian species with a 2-valved perianth. 


69. Horsfieldia carnosa Warb. Fig. 1C(69); 26 


Horsfieldia carnosa Warb., Mon. Myrist. (1897) 348, 619; Merr., En. Born. J. Str. Br. R. As. Soc. spec. 
number (1921) 268; Sinclair, Gard. Bull. Sing. 28 (1975) 21. — Myristica carnosa (Warb.) Boerl., 
Hand. Fl. Ned. Ind. 3, 1 (1900) 87 — Type: Beccari 344 (FI Acc. 7624), young male fl. (FI, n.v.); 
1242 (FI Acc. 7625), fr. (FI, n.v.; iso: K, P). 


Tree 4-10 m. Twigs terete, stoutish, towards apex 3-10(-16) mm diam., early 
glabrescent from a grey-brown tomentum composed of hairs c. 0.1 mm, bark of 
older twigs coarsely or finely striate, with a tendency to flake, drying somewhat 
pale, yellow-brown or light grey-brown, generally rather contrasting with the 
blackish brown colour of the dry petioles, lenticles usually not conspicuous. Leaves 
in 2 rows, chartaceous-coriaceous, rarely + membranous, elliptic-oblong to 
oblong, broadest at or slightly above the middle, 13-35 x 5-11 cm, base long- or 
sometimes short-attenuate, top acute-acuminate; upper surface drying bright dark 
brown, with a finely wrinkled-granulate structure, glabrous, lower surface early 
glabrescent (glabrous), without blackish dots; midrib above flat; nerves 13-18 pairs, 
flat above, the marginal arches indistinct; tertiary venation forming a lax network, 
faint or invisible on both surfaces; petioles 10-16 x 2-4 mm, glabrous or early 
glabrescent; leaf bud 9-13 x 2-3 mm, with grey-brown tomentum of hairs c. 0.1 mm 
long. Inflorescences densely to rather sparsely set with hairs c. 0.1 mm or less, in 
CO’: many-flowered, 3 or 4 times ramified, 6-17 x 5-14 cm, common peduncle 10-30 
mm; in 9: ramiflorous, rather many-flowered, c. 1-2 cm long, the flowers rather 
clustered; bracts elliptic to oblong, 4-10 mm long, pubescent, caducous. Flowers 
3-valved, glabrous, in CO in loose clusters of 3-9 each, pedicels 1-1.5 mm long, 
rather slender, glabrous, at base inarticulate. Male perianths globose or subglo- 
bose, 1.9-2.1 x 1.8-2.0 mm, top and base (broadly) rounded; pedicels 1.0-1.5 mm 
long; perianth at anthesis cleft to ¥3-/2, valves c. 0.2 mm thick, perianth not 
collapsing on drying. Androecium (sub)globose, 1.0-1.2 x 1.0-1.3 mm, circular in 
transverse section; anthers 9-11 (thecae 18-22), completely sessile (free apices + 0), 


Fig. 26. 


Horsfieldia carnosa Warb. 

a. portion of twig with leaf and male infloresces, x 1%; b. apical part of leafy twig, note pale 
colour of stem contrasting with dark drying colour of the petioles, x 2; c. mature male 
flower, lateral view, X 12; d. ditto, opened, showing androecium, x 12; e. androecium, 
longitudinal section, schematic, x 12; f. portion of twig with female inflorescence axillary to 
leaf scar, X 2; g. female flower in anthesis, lateral view, x 12; h. ditto, longitudinally 
opened, showing glabrous ovary with broadly 2-lipped stigmas, x 12; i. portion of older twig 
with infructescences, fruits mature, aril complete but torn by drying, x 2. —a. from van Niel 
5419; b-e. from S. 18011; f-h. from San. 63191; i. from San. 17438. 


223 


224 Gard. Bull. Sing. 38(2) (1985) 


curved, at apex concealing a small apical cavity 0.2-0.4 mm deep; column broad, + 
spongy, androphore rather narrow, distinct, 0.2-0.5 mm long, completely hidden 
by the anthers. Female perianth ellipsoid, c. 3.5 x 2.5 mm, glabrous, cleft toc. 4, 
valves c. 0.3-0.4 mm thick, pedicels c. 1-1.5 mm long, ovary ellipsoid, c. 2.0 x 1.5 
mm, glabrous, stigma consisting of two broad lips c. 0.2 mm high. Fruits 2-11 per 
infructescence, ellipsoid, top and base rounded, 1.6-2.0 x 1.2-1.5 cm, glabrous, 
drying brown with a finely granulated surface; pericarp c. 1.5 mm thick; stalk 1-2 
mm long; perianth not persisting. 


Distribution. Borneo: Sarawak, Brunei, Sabah, West Kalimantan (G. Klam). 


BORNEO. Sarawak: Beccari 1242; Clemens 22345; Haviland & Hose 2096; Native Coll. 676, 1970; 
S 15456, 15955, 18011, 32713: Sinclair & Kadim 10428 — Brunei: Fuchs et al. 21193; Van Niel 4519; 
Brtinig S. 1071 — Sabah: B.N.B. For Dept. 2378; SAN 17438, 63191, 73218, 80063 — West Kalimantan, 
G. Klam: H. Hallier 2381. 


Ecology. Heath forest, wet kerangas forest, peat swamp forest, Agathis- 
Casuarina forest; on white sandy soils; 0-100m alt; flowers mainly in July to 
November, fruits collected throughout the year. An extensive note on the ecology 
is given by Sinclair on p. 22. 


NOTES 


1. Fieldnotes. A low tree with slender trunk to 10 m high. Bark often flaking or 
shallowly fissured. Twigs light brown. Inner bark yellowish, thin, Sap watery, clear, 
not red or reddish. Sapwood whitish. Flowers greenish-yellow, anthers whitish. 
Fruits (immature) greenish-yellow, aril orange. 


2. This is a well-characterized species of low trees of the Kerangas or peat 
swamp forest, on white sand soils. It is distantly related to H. glabra, but easily 
distinguished by its usually stout habit, pale twigs with the bark tending to flake, 
the large and usually chartaceous-coriaceous (possibly somewhat fleshy) leaves, 
globose male flowers, the pedicels inarticulate at the base, the globose androecium 
with distinct but hidden androphore, and the small often + clustered fruits of c. 
1.6-2 cm length. 


70. Horsfieldia sterilis de Wilde, sp. nov. Fig. 1C(70) 


Folia membranacea, in sicco nigrescentia. Perianthium masculum late obovoideum. + carnosum, c. 1.7 
mm longum, 2-valvatum, alabastro in anthesi usque ad 14 fisso, androecio late obovoideo, c. 0.8 mm 
longo, apice non-excavato, antheris 3, c. 0.4 mm longis, sessilibus. — Type: Sabah, Aban Gibot SAN 
30597 (L; iso: K; SING, n.v.). 


Tree or shrub, 3-12 m. Twigs terete, not lined or winged, towards the apex 
2.5-4(-8) mm diam., dark brown or greyish brown, sometimes rather contrasting 
with the blackish colour of the dry petioles, bark early glabrescent, tomentum rusty 
to grey-brown, with hairs 0.1-0.2 mm; bark lower down finely striate, not tending 
to crack or flake; lenticels conspicuous or not. Leaves in 2 rows, membranous, 
elliptic-oblong to oblong(-lanceolate), broadest at or slightly above the middle, 
13-33 x 4.5-9 cm, base attenuate, top acute-acuminate; upper surface glabrous, 
drying rather dark brown to blackish, lower surface glabrous, drying brownish, 
without scattered larger dark-coloured dots; midrib flat above, glabrous; nerves 
11-17 pairs, flattish or slightly raised above, lateral arches not particularly distinct; 
tertiary venation forming a lax network, faint or invisible on both surfaces; petioles 
7-20 X 2-3.5 mm, glabrous (early glabrescent); leaf bud slender, 7-12 x 2-2.5 mm, 


New account of Horsfieldia 3 225 


densely (pale) rusty pubescent with hairs 0.1-0.2 mm long. Inflorescences sub- 
glabrescent, tomentum rather weak, of hairs 0.1-0.2 mm long, in CG: rather lax, 
10-20 x 5-8 cm with rather few side-branches, 2 or 3 times ramified, moderately 
densely flowered, common peduncle long, with several bract-scars, 30-90 mm long, 
flowers in loose clusters of 2-6 each; 9 inflorescences elongate, slightly-branched, 
almost spike-like, 5-10 cm long; perianths glabrous, 2-valved, pedicel glabrous, at 
base inarticulate; bracts ablong-lanceolate, 3-8 mm long, acute, pubescent with 
hairs 0.1-0.3 mm especially at the margins, caducous. Male perianth broadly 
obovoid-subglobose, 1.5-1.7 mm diam., top broadly rounded or somewhat depress- 
ed, base sub-attenuate, glabrous; pedicel 1-1.5 mm long, slender; perianth at 
anthesis cleft to c. ¥4, valves rather thick-fleshy, 0.3-0.4 mm thick, the perianth 
shrinking, not collapsing on drying. Androecium broadly obovoid, c. 0.8 X 0.6-0.7. 
mm, top + rounded, subcircular in transverse section; anthers 3 or (6?) (i.e. with 
5-6 thecae), completely sessile, c. 0.4 mm long, occupying only the apical part of 
the androecium, connectives broad, + triangular, with narrow thecae; column 
broad and solid, apical cavity absent or very inconspicuous; androphore conspi- 
cuous, broad, tapering to below, c. 0.4 mm long. Female flowers not seen; accord- 
ing to persistent perianths under immature fruits: perianth c. 2.5 mm long, 2- 
valved, glabrous. Fruits (immature ones, B.N.B. For. Dept. 27) 4-10 per spike-like 
infructescence, ellipsoid, top and base rounded, c. 2.2 X 1.8 cm, glabrous, drying 
blackish, not tubercled, pericarp c. 2 mm thick; perianth persisting under immature 
fruit; stalk 3-4 mm long. 


Distribution. Borneo: Sabah, a local endemic of SE. Sabah. 


BORNEO. Sabah, Tawau Dist.: (Orolfo) B.N.B. For. Dept. 27; (Aban Gibot) SAN 30597; (Shea) 
SAN 75748, 95959. 


Ecology. Smal! trees or shrubs in foresi on hiilsides and riverbanks; at. 80-500 m 
alt. Flowers April to July. 


Vernacular name. Duria (Sabah, Tawau Dist.}. 


NOTES 


1. Fieldnotes. Small trees or shrubs; bark greybrown, non-fissured. Inner bark 
with orange-red sap. Sapwood pale yellow. 


2. A species notable by its 2-valved perianth and the subconical broadly obovoid 
androecium of which only the apical half bears 3 or 6 rather much-reduced anthers; 
the basal part of the androecium or androphore is sterile, broad and tapering. 
Whether there are actually 6 anthers or only 3 anthers each with 2 thecae requires 
further confirmation. 


The structure of the androecium is reminiscent of that of species like H. crux- 
melitensis and H. clavata from New Guinea, but in these species the androecium Is 
much more elongate. In general habit the new species resembles e.g. H. pallidi- 
caula or H. sucosa, because of its rather pale twigs and blackish drying leaves, but 
our present species differs by much more elongated inflorescences. 


3. The type specimen was not treated under Horsfieldia by Sinclair. A second 
specimen (Orolfo, B.N.B. For. Dept. 27) was identified by him as H. bracteosa var. 
muicrocarya. 


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Comparative Anatomy of the Stipe of the Fern Genus 
Adiantum L. (Adiantaceae) 


'Aziz BIDIN & TREVOR WALKER 
‘Botany Department, Faculty of Life Sciences 
Universiti Kebangsaan Malaysia, Bangi, Selangor, Malaysia 
° Department of Plant Biology 
The University, Newcastle upon Tyne, NEI 7RU England 


Abstract 


The investigations carried out covering approximately half the species from virtually the entire 
geographical range of the genus Adiantum L. showed that there was a range of shape of the xylem 
strands proceeding by small steps from the simple (deep-crescent type) to the intricate form (tong- 
shaped). Eight types of xylem configurations were observed and the genus was subdivided accordingly. 


Introduction 


The fern genus Adiantum comprises about 150-200 species (Abraham et al., 
1962; Holttum, 1954; Tryon & Tryon, 1981) and has a nearly cosmopolitan dis- 
tribution except in regions with extremely dry or cold climates. The genus was 
subdivided into a number of groups based on its diverse morphological forms by 
early taxonomists such as Hooker & Baker (1874), Smith (1896), Christ (1897) and 
Diels (1902). Meanwhile Wylie (1948) investigated the role of leaf epidermis in 18 
species of Adiantum from America and Bidin (1980, 1984a, 1984b) looked at the 
cytology, leaf forms and leaf blade anatomy of the genus respectively. 


Anatomical and morphological studies of the stipe have also been regarded as 
useful methods in solving taxonomic problems of ferns. Pres! (1847) and Thomas 
(1886) investigated the structure and arrangement of vascular bundles in the stipes 
of a wide range of ferns and drew attention to the great variation found. Later 
authors used these characters in investigating particular taxonomic problems, e.g., 
Milde (1866) in distinguishing between Athyrium and Asplenium, whilst Ching 
(1940) used them in the subdivision of Polypodiaceae s.l. More recently Keating 
(1968) studied the stipe anatomy of the Dennstaedtioid genera, while Lucansky & 
White (1974) made comparative studies of the nodal and vascular anatomy of 
neotropical members of Cyatheaceae. In Taiwan, Lin & DeVol (1977) prepared a 
multiple choice key based on the study of stipe characters to the species, genera and 
families of local ferns. 


Materials and Methods 


Plant material for the study were gathered from various geographical areas by 
the authors. Live plants were grown at the Moorbank Experimental Garden, 
Newcastle University, whilst dried specimens were kept at the Herbarium of the 
Plant Biology Department of the same University. 


Segments were cut from three positions along the stipe; i.e., at the base, in the 
middle region and the upper portion just below the first rachis or pinnule. The 
segments were each about | cm long. Fresh material was fixed in 70% alcohol. 
Extremely hard stipes and those obtained from the herbarium specimens were 


227 


228 Gard. Bull. Sing. 38(2) (1985) 


softened by boiling in water for two hours prior to fixation. Various methods of 
sectioning were employed. In most cases sections were obtained by free hand or by 
use of the sliding microtome but for hard materials it was necessary to embed in 
paraffin wax before microtoming. Staining in all cases was in Safranin 0 and 
Light/Fast Green. 


Results and Discussion 


The stipe of Adiantum is usually dark brown to black, slender and has a 
polished-glossy appearance. Some species are however sparsely hairy or scaly 
especially towards the base. 


The anatomical structure of the stipe is fundamentally similar to that of the 
rhizome. It possesses a layer of longitudinally elongated epidermal cells, a cortex, 
an endodermis and a stele. In all cases the endodermis surrounds each vascular 
bundle and in certain species the stele is clearly visible in a cross-section even with 
the unaided eye. The stelar system in the stipe of Adiantum consists of one or two 
traces which are usually arranged in an adaxially curved arc and shows a wide range 
of variation. The stele in cross-section may have a terete or undulate outline or 
sometimes an intermediate form. In some species the outlines of the endodermis 
bear no specific relation to the cross-sectional shape of the stipe (Table 1). The 
arrangement of strands may change gradually from base to apex, and all the 
changes however minor should be noted, particularly those affecting the xylem. 


Two bundles enter the base of the stipe, and later join upwards to form a 
four-angled strand. This can be clearly seen in the majority of the species. In some 
cases the double nature of the bundle is not just confined to the basal region but 
many extend upward to the middle or even the upper region of the stipe. Observa- 
tions show that even the double nature of the bundle is not uniform throughout the 
genus. In the Reniforme and Caudatum groups there is only one bundle present 
even at the extreme base of the stipe and it persists up the entire length. 


Except for the species showing an arc-shaped xylem throughout the entire length 
of the stipe, all members of Adiantum show two Onoclea-type strands in the basal 
region. Each of these strands contains a hippocampus-shaped xylem mass with 
hooked or blunt or even sharp ends towards the groove of the stipe. The protox- 
ylem occurs on the inner surface, and a phloem layer surrounds the xylem mass. In 
some cases the two bundles are more widely separated towards the groove (adaxial) 
side of the stipe than at the opposite rounded (abaxial) side. Further up the stipe 
their abaxial ends come in contact and fuse to form a V-shaped, basin-shaped or 
tongs-shaped bundle with ends which differ in detail from one to another. 


A more detailed examination of a large number of taxa (5Q) reveals that it 1s not 
only basin-shaped or V-shaped xylem strands which can be found in Onoclea-type 
meristeles in the upper region o the stipe (Plate 1). The shape develops gradually 
from a simple crescent configuration in the Caudatum and Reniforme groups (Fig. 
1) to a more complex one in the shape of tongs as in A. /ucidum and A. tetraphyl- 
lum (Fig. 8). The first sign of change is that the crescent-shaped strand begins to 
show a weak constriction in the middle which finally divides the strand into two 
halves. The tips of the halves curve downwards and are blunt with no hooks present 
(Fig. 3). This is a transitional shape between the simple crescent-like and the more 
complex, tongs-like configuration and is found in A. /unulatum. 


FP, 


Fig. 1: Deep crescent Fig. 2: Light crescent Fig. 3: Bird-shaped 


ww wy’ \/ 


Fig. 4: Slightly curved- Fig. 5: Saucer-shaped 
upward Fig. 6: V-shaped 


Fig. 7: Basin-shaped Fig. 8: Tongs-shaped 


Plate 1. Shape of xylem strands in the stipe of Adiantum. The types are arranged in order of increasing 
complexity, which is believed to coincide with the direction of evolutionary change. The 
numbers in this figure correspond to the number of the groups in Table | of the types of supe 
in Adiantum. 


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The next type of xylem strand is still simple in form but in this case the tips of the 
two halves point upwards and are blunt and not hooked (Fig. 4). This shape is 
found in A. formosum and all members of the Capillus-Veneris group. The strands 
in A. patens is a modification of this; here the tips of the two halves are sharp and 
curve upwards, whilst the centre of the strand makes a shallow curve downwards 
forming a saucer-shaped strand (Fig. 5). A further development may be seen in A. 
hispidulum, A. cunninghamii, A. affine,- A. whitianum, A. silvaticum and A. 
pedatum. In these examples the xylem strand has a very characteristic V-shaped 
configuration and the tips of the two halves are sharp and a little curved (Fig. 6). 


In some species such as A. macrophyllum and A. serratocristatum where the 
fluting of the stipe is very deep the abaxial arms of xylem join to form a basin- 
shaped strand with a nearly flat base. The arms are very slender and well separated 
from each other in the central region (Fig. 7). In other cases the metaxylem 
portions although fusing with each other fail to develop fully in the central region of 
the abaxial part of the strand resulting in a shape similar to a pair of tongs with long 
curving handles (Fig. 8). This type is found in almost all members of the Polysorus 


group. 


Conclusion 


From the observations, it will be seen that there 1s a sequence of shapes of the 
xylem strands ranging by small steps from the deep-crescent type seen in A. 
reniforme to the rather intricate tongs-shaped type seen in A. tetraphyllum. This 
sequence is illustrated in Plate 1. Part of the sequence was described by Ogura 
(1972) but with important gaps present. 


Lin & De Vol (1977) in their investigation of the ferns of Taiwan stated in the key 
to the identification of genera that Adiantum had a V-shaped type of xylem 
configuration. There is no indication of whether or not more than one species was 
examined and indeed no direct indication of what species it was. However Tsai 
(1972 and 1973) lists 53 ferns of Taiwan which he has cytologically examined and 
the only species figuring in the list is A. capillus-veneris. This species has indeed a 
shallowly V-shaped xylem configuration. It is evident that a key of this nature 
based on inadequate samplings would not necessarily work in other regions and 
such information may be misleading if indiscriminately applied. 


Acknowledgements 


This work is partially supported by Universiti Kebangsaan Malaysia Research 
Grant No. 18/82 for which we are very grateful. 


References 


Abraham, A., C.A. Ninan and P.M. Mathew, (1962). Studies on the cytology and 
phylogeny of the pteridophytes. J. Ind. Bot. Soc., XLI (3): 339-421. 

Bidin, A. (1980). Studies in the fern genus Adiantum L. Ph.D. Thesis, University 
of Newcastle upon Tyne. (Unpublished). 

Bidin, A. (1984a). The importance of leaf forms in the subdivision of the genus 
Adiantum L. Sains Malaysiana, 13(3): 279-289. 


EE 


Comparative stipe anatomy of Adiantum 233 


Bidin, A. (1984b). Leaf blade anatomy of some species of Adiantum L. Sains 
Malaysiana, 13(3): 291-301. 

Ching, R.C. (1940). On natural classification of the family Polypodiaceae. Sunyat- 
senia, 5(4): 201-268. 

Christ, H. (1897). Die Farnkrauter der Erde. Jena. 

Diels, L. (1902), in Engler, A. and K. Prantl, Die natiirlichen Pflanzenfamilien, 
Leipzig. 

Holts R.E. (1954). Flora of Malaya, Vol. II. Ferns of Malaya, Singapore. 

Hooker, W.J. and J.G. Baker (1874). Synopsis Filicum 2nd Ed., London. 

Keating, R.C. (1968). Trends of specialisation in the stipe anatomy of Dennstaedtia 
and related genera. Am. Fern J., 58: 126-139. 

Lucansky, T.W., and R.A. White (1974). Comparative studies of the nodal and 
vascular anatomy in the neotropical Cyatheaceae, III. Nodal and petiole pat- 
terns; summary and conclusions. Am. J. Bot., 61(8): 818-828. 

Lin, B.L., and C.E. DeVol (1977). The use of stipe characters in fern taxonomy, I. 
Taiwania, 22: 91-99. 

Milde, J. (1866), Das Genus Athyrium. Bot. Zeito, 24: 373-376. 

Ogura, Y. (1972). Comparative anatomy of vegetative organs of the pteridophytes. 
2nd. Ed. Gebruder Borntraeger. Berlin, Stuttgart. 

Presl, K.B. (1847). Die Gefassbiindel in Stipes der Farn. Abh. Konigl. Bohm. 

Gesell. Wissen.: 1-48. 

Smith, J. (1896). Ferns: British and foreign. London. 

Thomae, K. (1886). Die Blattstiele der Farne. Ein Beitrag zur bergleichenden 
Anatomie. Jb. Wiss. Bot. 17: 99-161. 

Tryon, R.M. and A.F. Tryon (1981). Ferns and allied plants. Springer Verlag, New 
York, Heidelberg. Berlin. 

Tsai, J.L. (1972). Chromosome numbers of some Formosan ferns (1). Jour. Science 
and Engineering, 9: 125-132. 

Tsai, J.L. (1973). Chromosome numbers of some Formosan ferns (2). Jour. Science 
and Engineering 10: 261-275. 

Wylie, R.B. (1948). The dominant role of the epidermis in leaves of Adiantum. 
Am. J. Bot. 35: 465-472. 


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